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1 4 MAR /983 


Bulletin of the 


British Museum (Natural History 

Zoology series Vol 42 1982 

British Museum (Natural History) 
London 1983 

Dates of publication of the parts 

No 1 25 March 1982 

No 2 29 April 1982 

No 3 27 May 1982 

No 4 24 June 1982 

ISSN 0007-1498 

Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset 

Zoology Volume 42 


No 1 The tick collection (Acarina: Ixodoidea) of the Hon. Nathaniel 
Charles Rothschild deposited in the Nuttall and general collections of 
the British Museum (Natural History) 
By James E. Keirans 1 

No 2 Hydroids and medusae of the family Campanulariidae recorded from 
the eastern North Atlantic, with a world synopsis of genera 
By P. F. S. Cornelius 37 

No 3 Miscellanea 

A new species of free-living nematode from the Firth of Clyde, Scotland 

By P. J. D. Lambshead 149 

Notes on Atlantic Asteroidea 2. Luidiidae 

By A. M. Clark 157 

New and little known species of Oncaeidae (Cyclopoida) from the 

Northeastern Atlantic 

By S. J. Malt 185 

Larval and post-larval development of the Slender-legged Spider Crab, 
Macropodia rostrata (Linnaeus), (Oxyrhyncha: Majidae: Inachinae), 
reared in the laboratory 
By R. W. Ingle 207 

No 4 Miscellanea 

New species of marine nematodes from Loch Ewe, Scotland 

By H. M. Platt and Z. N. Zhang 227 

The larval development of Crangon crangon (Fabr. 1 795) (Crustacea : 


By A. R. Gurney 247 

A revision of the spider genus Cocalodes with a description of a new 

related genus (Araneae: Salticidae) 

By F. R. Wanless 263 

Anatomy and evolution of the jaws in the semiplotine carps with a 
review of the Genus Cyprinion Heckel, 1843 (Teleostei: Cyprinidae) 
By G. J. Howes .299 

Bulletin of the 

British Museum (Natural History) 

The tick collection (Acarina: Ixodoidea) 
of the Hon. Nathaniel Charles Rothschild 
deposited in the Nuttall and general 
collections of the British Museum 
(Natural History) 

James E. Keirans 

With a Foreword by the Hon. Miriam Rothschild 

Zoology series Vol 42 No 1 25 March 1982 

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
ever-growing collections of the Museum, both by the scientific staff of the Museum and by 
specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
works of reference that will remain indispensable for years to come. 

Parts are published at irregular intervals as they become ready, each is complete in itself, 
available separately, and individually priced. Volumes contain about 300 pages and several 
volumes may appear within a calendar year. Subscriptions may be placed for one or more of 
the series on either an Annual or Per Volume basis. Prices vary according to the contents of 
the individual parts. Orders and enquiries should be sent to: 

Publications Sales, 

British Museum (Natural History), 
Cromwell Road, 

London SW7 5BD, 

World List abbreviation: Bull. Br. Mus, not. Hist. (Zool.) 

Trustees of the British Museum (Natural History), 1982 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology : Dr J. G. Sheals 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor : Mr C. G. Ogden 

ISSN 0007-1498 Zoology series 

Vol 42 No 1 pp 1-36 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD Issued 25 March 1982 


The tick collection ( Acarina: Ixodoidea) of thV ^ 1982 

\ -4 LIBRARY "^ 

Hon. Nathaniel Charles Rothschild deposited in ^zy 
the Nuttall and general collections of the BritisBT 
Museum (Natural History) 

James E. Keirans 

Department of Health and Human Services, Public Health Service, National Institutes of 
Health, National Institute of Allergy and Infectious Diseases, Rocky Mountain Laboratory, 
Hamilton, Montana 59840, U.S.A. 

With a Foreword by the Hon. Miriam Rothschild 


Synopsis 1 

Foreword 3 

Introduction 4 

PART I Rothschild specimens donated to Professor G. H. F. Nuttall ... 5 
Species of Ixodoidea represented in the Nuttall collection, British Museum 

(Natural History), donated by the Hon. N. C. Rothschild .... 5 

Rothschild specimens in the Nuttall collection with type status ... 6 

Rothschild specimens in the Nuttall collection 6 

A single tick collection given by the Hon. Walter Rothschild to 

G. H. F. Nuttal via Guy A. K. Marshall 17 

Rothschild specimens missing from Nuttall collection 17 

PART II Rothschild specimens donated to British Museum (Natural History). 20 
Species of Ixodoidea represented in the collection of the British Museum (Natural 

History), donated by the Hon. N. C. Rothschild 20 

Rothschild specimens in the British Museum (Natural History) collection with 

type status 21 

Rothschild specimens in British Museum (Natural History) collection . . 22 

Rothschild specimens missing from British Museum (Natural History) collection 32 

Locality list for ticks in the Rothschild collection 32 

Host list for ticks in the Rothschild collection 34 

Acknowledgements 35 

References 36 


The Hon. N. C. Rothschild achieved worldwide fame for his study of the Siphonaptera. 

During the early part of this century he also made a significant collection of ticks which he 
gave to Professor George H. F. Nuttall at Cambridge University. Upon Nuttall's death, the 
collection was given to the British Museum (Natural History) and is kept as a separate entity. 

Bull. Br. Mus. nat. Hist. (Zool.) 42( 1 ): 1 -36 Issued 25 March 1 982 


Male Ixodes uriae White, 1852. (Ixodoidea: Ixodidae). A tick with a circumpolar distribution in 
both the northern and southern hemispheres. It is ectoparasitic on a variety of seabirds and to 
date, twenty different arboviruses have been recovered from this species. 


In addition, Rothschild on numerous occasions between 1911 and 1923 deposited ticks in 
the British Museum (Natural History). Herewith are recorded the 197 tick collections of the 
Hon. N. C. Rothschild plus one of the Hon. Walter Rothschild deposited in the Nuttall 
collection and the 156 collections in the general collection of the British Museum (Natural 


In 1913 N. C. Rothschild presented his collection of 'Siphonaptera and other parasitic 
insects' to the British Museum (Natural History). One of the conditions attached to the gift 
read as follows: 'The Trustees shall so soon as the said collection comes into their hands or is 
placed under their control and as soon as the funds placed at their disposal by Parliament 
will allow cause to be made and published a catalogue giving the names and full number of 
specimens of each species of parasitic insect contained in the said collection.' 

Over 60 years has now elapsed since the collection was accepted, but the catalogue has not 
yet been completed. Although ticks are arthropods, they are not insects, but this group of 
animals, along with a number of mites, was included in the collection. 

During my period of Trusteeship of the British Museum (Natural History) I had the 
opportunity of discussing this problem with Harry Hoogstraal, and he suggested that James 
Keirans might undertake to catalogue the Rothschild ticks. To my great delight he agreed to 
do this, although the task proved more time-consuming than we had at first anticipated. This 
was because the Rothschild Collection had not been preserved as a single unit, and the ticks, 
along with the Nycteribiidae, Cimicidae, Hippoboscidae and other smaller groups, had been 
distributed in the general collection and, in addition, had not all received Museum accession 

Although these circumstances added enormously to James Keirans' labours, they made 
the compilation of this Catalogue even more necessary and valuable. Until those who are 
engaged in ecological and medical research come to use museum material such as this, no 
one can really appreciate the immense boon of well-catalogued individual collections. 

Since, between 1900 and 1910, N. C. Rothschild had given Professor Nuttall the bulk of 
his tick material*, it was considered more appropriate to merge the two catalogues into one, 
and the manuscript was therefore divided into two parts: I, the specimens collected by 
Rothschild, now in the Nuttall collection; and II, the specimens donated by Rothschild to 
the British Museum (Natural History). The ticks in the general collection have been 
added a course we followed when cataloguing the fleas (G. H. E. Hopkins & M. Rothschild, 
Vols. I-V, 1953-1 971) and the Nycteribiidae (Oscar Theodor, 1967). 

It will be noted that Walter Rothschild contributed one specimen to this collection. It has 
not usually been appreciated that N. C. Rothschild's brother also added considerably to the 
flea material, although he did not describe any species himself. The N. C. Rothschild 
donations of Nycteribiidae to the British Museum contained 56 undescribed species, and the 
Ixodoidea collections 10. The two brothers and their collaborators described over 5000 new 
species, but they probably collected twice that number of animals new to science. 

James Keirans has added greatly to the value of this compilation by providing information 
concerning the viruses and in certain cases rickettsiae of which the Rothschild ticks are 
known vectors. 

The illustration depicts a male specimen of Ixodes uriae White, one of the most 
spectacular species, adorned with a handsome fringe along the posterior margin, which has 
the dubious distinction of carrying at least 20 known viruses in five different serogroups. 

*After the sudden death of Professor Nuttall in December 1937 his collection was presented 19 the British Museum 
(Natural History) by the Molteno Institute. In accordance with the terms of the gift this collection is maintained as a 
discrete entity. 


Altogether the meticulous care and accuracy with which James Keirans has completed 
this catalogue is beyond praise. 

Miriam Rothschild 


Early in the twentieth century, Nathanial Charles Rothschild acquired a large tick 
collection, primarily from collectors who, knowing of his great interest in fleas, would send 
him not only that group but also other ectoparasites including flies, bugs, and ticks. 

He deposited his tick collection in two institutions; the Molteno Institute for Research in 
Parasitology, Cambridge University, where Professor G. H. F. Nuttall made the original 
determinations of all tick material, and the British Museum (Natural History), where Mr A. 
S. Hirst made the tick determinations. As mentioned earlier, the Nuttall collection is now at 
the BM(NH). 

Because the Nuttall collection is kept as a discrete entity, separate from but housed with 
the BM(NH) collection, parts of Rothschild's tick collection are in each of the above 
collections. This catalogue is also divided into two sections: Part I, Rothschild tick donations 
to the Nuttall collection; Part II, Rothschild tick donations to the BM(NH). 

The systems for numbering tick collections are different for the two sections. In Part I, the 
number used is that given by Professor Nuttall to a collection. The first entry is N (Nuttall) 
72 followed by N73, N74, etc., in ascending sequence, first for existing specimens, then for 
specimens which are now missing. Each Nuttall number is followed by a Rothschild 
collection number or the word 'None' if the collection received no Rothschild number. A 
Nuttall number refers to a handwritten entry in Professor Nuttall's tick catalogue, which is 
now kept in the Arachnida and Myriapoda Section of the BM(NH). The locality and host 
data are cited in their modern forms, but where these could not be established Nuttall's own 
entries are quoted. 

One additional point should be made relating to the Nuttall collection forming Part I. 
Nathaniel C. Rothschild donated all specimens recorded in Part I with the single exception 
of N888 which was donated by his brother, the Hon. Walter Rothschild, to the Entomo- 
logical Research Committee and subsequently to Professor Nuttall. 

Collections in Part II are entered alphabetically by genus, first Argasidae, then Ixodidae. 
All information is presented as in Part I with the exception that replacing a Nuttall and 
Rothschild number will be a British Museum (Natural History) registration number. Where 
no registration number was given to a collection, the word 'None' will appear. I have 
given each collection a set of Keirans numbers, K1-K156. These numbers, along 
with the Nuttall numbers from Part I, will allow the reader to use the tick species, 
locality, and host lists and refer back to individual collections within Parts I and II. 

Throughout the text, citations are given to publications which mention a particular taxon. 
My comments are presented within square brackets [ ]; those comments by Nuttall or some 
other investigator are cited within square brackets enclosed by quotation marks '[ ]'. 



Rothschild specimens donated to the Nuttall collection 

Species of Ixodoidea represented in the Nuttall collection, British Museum 
(Natural History), donated by the Hon. N. C. Rothschild 

Nuttall numbers follow each species. Numbers in parentheses refer to missing collections. 
Thus, validity of original determinations is unverified. 


Argas boueti Robaud and Colas-Belcour 1 164 
Argas persicus (Oken) 589, 590, (3536)) 
Argas reflexus (Fabricius) (3 1 59) 
Argas vespertilionis (Latreille) 591, 592, 593, 

1151,1152,1153, 1163 
Argas sp. 11 54, (1388) 

Ornithodoros capensis Neumann group 594 
Ornithodoros moubata (Murray) 595 

Otobius megnini (Duges) (596) 


Amblyomma albolimbatum Neumann (78), 567, 


Amblyomma australiense Neumann 547 
Amblyomma cajennense (Fabricius) 557, 648 % 
Amblyomma clypeolatum Neumann (75) 
Amblyomma compressum Macalister 564 
Amblyomma cyprium Neumann 73, 565 
Amblyomma decoratumC. L. Koch (93) 
Amblyomma dissimile C. L. Koch 568 
Amblyomma gemma Donitz 558b 
Amblyomma geoemydae (Cantor) 545 
Amblyomma hirtum Neumann (397) 
Amblyomma longirostre(C. L. Koch) 560 
Amblyomma marmoreum C. L. Koch (77), 570 
Amblyomma ovaleC. L. Koch 562, 566 
Amblyomma pecarium Dunn 557 
Amblyomma tholloni Neumann 559 
Amblyomma triguttatum C. L. Koch 546, (551), 


Amblyomma varium C. L. Koch (3351) 
Amblyomma sp. 74, (76), (544), 633, 1 166, 1386, 

1391, 1392, 1393, 1394, 1396, 1397 

Aponomma decorosum (L. Koch) 550 
Aponomma gervaisi (Lucas) (92), (548) 
Aponomma gervaisi var. lucasi (Warburton) (90), 


Aponomma hydrosauri (Denny) 556 
Aponomma varanensis (Supino) 554 
Aponomma sp. (57 1 ), 3 1 38 

Boophilus decoloratus (C. L. Koch) (3157) 
Boophilus microplus (G. Canestrini) 588 

Dermacentor albipictus (Packard) 666, 1105, 

1497, 1498, 1499, 1500, 1501, 1502, 1503 
Dermacentor andersoni Stiles 1387, 1390,3502 
Dermacentor reticulatus (Fabricius) 3 1 60 
Dermacentor rhinocerinus (Denny) 555, 563 
Dermacentor sp. 3330 

Haemaphysalis bispinosa Neumann 574, 582, 


Haemaphysalis campanulata Warburton 579 
Haemaphysalis celebensis Hoogstraal, Trapido 

and Kohls 72 

Haemaphysalis elongata Neumann 670 
Haemaphysalis erinacei Pavesi 3537, 3538, 3539 
Haemaphysalis humerosa Warburton and Nuttall 

Haemaphysalis hystricis Supino 572, 573, 575, 


Haemaphysalis indica Warburton 575x 
Haemaphysalis leachi (Audouin) (539), 577, 578 
Haemaphysalis punctata G. Canestrini and 

Fanzago 580 

Haemaphysalis tiptoni Hoogstraal 670 
Haemaphysalis sp. 58 1 

Hyalomma aegyptium (Linnaeus) 549, 56 1 
Hvalomma rufipes C. L. Koch 558a, 888 (W. 

Hyalomma syriacum C. L. Koch (3139), (3140), 


Hyalomma truncatum C. L. Koch 1 167 
Hyalomma sp. 1398, (3 136) 

Ixodes australiensis Neumann 645, 646 

Ixodes boliviensis Neumann 637 

Ixodes brunneus C. L. Koch (396) 

Ixodes cookei Packard 1 504 

Ixodes fecialis Warburton and Nuttall 650, 1211, 


Ixodesfrontalis (Panzer) 523 
Ixodes hexagonus Leach (216), (217), (298), 

(322), (324), 597, 598, 599, 600, 601 , 602, 603, 

604, 606, 607, 608, 609, 610, 611, 612, 613, 

(674), (746), 1067, 1159, 1160 
Ixodes holocyclus Neumann 643, (644), 1212 
Ixodes lividus C. L. Koch 605, 1068 


Ixodes loricatus Neumann 638, 639, 640, 64 1 

Ixodes luciae Senevet 647 

Ixodes nitens Neumann 360 

Ixodes putus (O. Pickard-Cambridge) (516), 

(5 17), (5 18) 
Ixodes ricinus (Linnaeus) (509), (628), (629), 

(630), (631), (632), (656) 
Ixodes rothschildi Nuttall and Warburton 634 
Ixodes rubidus Neumann 826 
Ixodes scapularis Say 626 
Ixodes texanus Banks 1 399 
Ixodes trianguliceps Birula 619, (620), 621, 622, 

623,624,625,1069, 1161 

Ixodes unicavatus Neumann (395), 652, 667 
Ixodes uriae White 617,618 
Ixodes vespertilionis C. L. Koch 649 
Ixodes vestitus Neumann 642 
Ixodes sp. 1066, 1 162, 1 167, 1389 

Rhipicephalus evertsi Neumann (3 1 57) 
Rhipicephalus haemaphysaloides Supino 584, 


Rhipicephalus muehlensi Zumpt 585 
Rhipicephalus sanguineus (Latreille) (586), (587) 
Rhipicephalus sp. 1 395 

Rothschild specimens in the Nuttall collection with type status 








Haemaphysalis celebensis Hoogstraal, Trapido, and Kohls, 1965. J. 

Paras it. 51: 1001, figs. 1-9. 

Ixodes nitens Neumann, 1904. Archs. Parasit. 8: 459. 

Amblyomma quasicyprium Robinson, 1926. Ticks. A Monograph of the 

Ixodoidea Pt. 4: 237, fig. 1 17. 

Ixodes percavatus rothschildi Nuttall and Warburton, 1911. Ticks. A 

Monograph of the Ixodoidea Pt. 2: 22 1 . 

Ixodes loricatus spinosus Nuttall, 1910. Parasitology, Cambridge 3: 

411, fig. 5. 

Ixodes fecialis Warburton and Nuttall, 1909. Parasitology, Cambridge 

2: 58, figs. 1-2. 

Haemaphysalis humerosa Warburton and Nuttall, 1909. Parasitology, 

Cambridge 2: 60, figs. 4-5. 

Rothschild specimens in the Nuttall collection 

N72 R916 

1 9 Haemaphysalis hystricis (Haemaphysalis 

celebensis HOLOTYPE) 
ex. Sus celebensis (Sus verrucosus celebensis) 
Celebes. No date (Sulawesi 02WS, 121WE, 


Presented as mounted specimen No. 916 which 
we unmounted 4.1. 1 9 1 5, C. W. [Cecil Warburton] 
det. 8.1.1915. 

Publications: Nuttall & Warburton (1915: 425, 
426); Hoogstraal el al. (19656: 1001); Hoogstraal 
etal. (1973: 556). 

N73 R915 

1 d Amblyomma caelaturum [This species epithet 

crossed out and cyprium inserted] 

(Amblyomma cyprium) 

ex. Sus celebensis (Sus verrucosus subsp.) 
Celebes. No date (Sulawesi 02WS, 12100'E, 


Presented as slide No. 9 1 5 which was unmounted, 
being defective 5.1.1915. 
3 d 1 Amblyomma cyprium mounted as slides, 2 

transparent, 1 opaque see slides No. 73 (i) (ii) 

(iii) (Nos. 917, 919, 920) (3 slides) (L. E. 

Robinson det. 26.1. 1915) redet. 26.111. 1919. 
Note: The 3 slide-mounted males were missing 
from the Nuttall collection in 1977. 
Publication: Robinson (1926: 236). 

N74 R918 

1 N Amblyomma (? caelaturum Cooper & 
Robinson) (Amblyomma sp.). Unmounted 


from slide labelled as off Sus celebensis (Sus 

verrucosus subsp.) 
Celebes. No date (Sulawesi 02WS, 121WE, 

(L. E. Robinson det. 26.1. 1915). 

N360 None 

1 9 Ixodes nitens COTYPE (Ixodes nitens 


ex. Mus macleari (Rattus macleari) 
Christmas Island, Pacific Ocean (1039'S, 


(ex. Shipley coll. No. 2) 

Publications: Neumann (1904: 460); Nuttall & 
Warburton(1911: 185). 

N523 None 

1 9 Ixodes brunneus (minus hypostome, digits) 

(Ixodes fron tails) 
ex. Passer montanus 
Saint-Genies-de Malgoires (4357'N, 0413'E), 

Card, France 
16.XI. 1908 Albert Hugues 
Publication: Nuttall & Warburton (1911:1 92). 

N545 R258 

Amblyomma sp. [sex/stage not given] (1 9 
Amblyomma geoemydae) (tentative deter- 

Found on ground at Paquil, Luzon, Philippine 
Islands (locality not verified) 

15.1.1895 A. Everett 

Publication: Robinson (1926: 48) recorded thfs as 

A. americanum. 

N546 R174 

Amblyomma triguttatum [sex/stage not given] (4 9 

Amblyomma triguttatum) 
ex. kangaroo 
Barrow Island (2048'S, 115'23'E), N. W. 


XI. 1900 C. J. T. Tanney. det. L. E. Robinson 
Publication: Robinson (1926: 57) indicated d's 
present but not in Nuttall collection in 1977. 

N547 R274 

6 d, 1 9 Amblyomma australiense 

ex. Echidna aculeata(Tachvglossus aculeatus] 

Mt Anderson ( 1 758'S, 1 24 ; 04'E), W. Australia 

[No date] C. J. T. Tanney 

Publications: Robinson (1926: 135, 136) 

illustrated d and 9 from this lot. Taylor (1946: 

100) repeated Robinson's descriptions and 


N549 R254 

Hyalomma syriacum [sex/stage not given] (4 cf 

Hyalomma aegyptium) 
ex. tortoise 
No data 
Note: An additional 1 d, 1 9 of this collection sent 

to New York National Museum, New York 
City, 18.IV.1913. 

N550 R196 

Aponomma decorosum [sex/stage not given] (2 d 

Aponomma decorosum) 
ex. Australian monitor 

Note: A. decorosum is found only in Australia and 
ranges from Queensland south to Victoria. 

N552 R179 

Amblyomma triguttatum [sex/stage not given] (2 

d, 3 9 Amblyomma triguttatum) 
ex. kangaroo 
Barrow Island (2048'S, 115'23'E), N. W. 


20.XI.1900C.J. T. Tanney 
Publication: Robinson (1926: 57). 

N553 R211 

Amblyomma triguttatum [sex/stage not given] (4 

9, 13 N Amblyomma triguttatum) 
ex. kangaroo 
Barrow Island (2048'S, 115'23'E), N. W. 


18.XI.1 900 C.J.T. Tanney 
Publication: Robinson (1926: 57). 

N554 R236 

Aponomma exornatum [sex/stage not given] (2 d 

Aponomma varanensis) 
ex. Monitor strix [name not verified] 
? Australia 

Note: Collection probably not from Australia. 
The only collection of A. varanensis from 
Australia is the type d of A. quadratus (an 
inornate form of A. varenensis). This single 
record may have been an accidental importation 
of tick and host. 

N555 R259 

Dermacentor rhinocerotis [sex/stage not given] (7 

d Dermacentor rhinocerinus) 
No other data 

Note: There is an additional d of this species 
mounted on a slide. 

N556 R287 

Aponomma hydrosauri [sex/stage not given] (2 d, 

1 9 Aponomma hydrosauri) 
? Adelaide (3455'S, 1 3835'E), W. Australia 

N557 R198 

Amblyomma cajennense [sex/stage not given] (1 

d, 2 9 Amblyomma cajennense; 3 d 

Amblyomma pecarium) 
ex. peccary Tayassu sp. 
Santa Andrea, Tabasco 
30. V.? '[No particulars]' 

Note: The above locality information was taken 
from the vial label. It should be San Andres, 
Estado de Tabasco ( 1 8WN, 9240'W), Mexico. 



Publication: Robinson (1926: 53) incorrectly gave 
the Nuttall number as 577. 
Amblyomma cajennense is a vector of Wad 
Medani virus and the rickettsia of Rocky 
Mountain spotted fever. 

N558a R263 

1 d, 1 9 Hyalomma aegyptium (Hyalomma 

ex. giraffe, Giraffa camelopardalis 

No other data 

Hyalomma rufipes is the vector of the viruses 

Dugbe, Tete, Matruh, and Crimean Congo 

hemorrhagic fever. 

N558b R263 

2 d Amblyomma gemma 

ex. giraffe, Giraffa camelopardalis 

No other data. 

Amblyomma gemma is a vector of Nairobi 

sheep disease virus. 

N559 R285 

1 9 Amblyomma tholloni 

No host 

Mt Ruwenzori (0023'N, 2954'E), on the 

Uganda-Zaire border, Africa 1906 A. F. R. 


N560 R260 

1 d Amblyomma longirostre 

ex. Coendou simonsi (Coendou bicolor simonsi) 

Charapaya(1722'S, 6645'W), Bolivia 

22.VI.1901 P.O.Simons 

Publication: Robinson (1926: 140). 

N561 None 

Hyalomma syriacum [sex/stage not given] (4 d 

Hyalomma aegyptium) 

ex. Algerian tortoise 

N. C. Rothschild 

N562 R261 

Amblyomma ovale [sex/stage not given] (2 d 

Amblyomma ovale) 
ex. Speothas venaticus 
Charapaya(1722'S,66'45'W), Bolivia 
27.VI.1901 P.O.Simons 

N563 R266 

10 d '[small and large]' Dermacentor rhinocerotis 
(7 cf Dermacentor rhinocerinus) 
ex. rhinoceros 
No other data 

Note: Nuttall, in his notebook containing infor- 
mation on ticks sent to various individuals and 
institutions, indicated that this collection was 
from Nyasaland, British Central Africa [Malawi] 
in 1907. One additional d of this collection was 
sent to Dr J. G. Parham, Marromeu, Zembezia, 
Portuguese East Africa [People's Republic of 
Mozambique], 18.IV.1913 and one additional d 

was sent to New York National Museum, New 
York City, 18.IV.1913. 

N564 R218 

Amblyomma badium [The species epithet crossed 

out and sublaeve inserted] (6 d Amblyomma 


ex. white-bellied pangolin, Manis tricuspis 
No other data 
L. E. Robinson det. 1926 

N565 R243 

1 9 Amblyomma quasicyprium Robinson, 1926 

TYPE (Amblyomma cyprium) 
ex. Spider monkey Ateles melanochoerus [This 

species epithet is crossed out and melanochir 

inserted] (Ateles g. geoffroyi) 
Frontera, ( 1 832'N, 9238'W), Tabasco, Mexico 
Notes: Both host and locality are incorrect for 
Amblyomma cyprium although this 9 appears to 
be true A. cyprium. 

Nuttall 575 and 3330 also bear Rothschild 
number R243. 

Publications: Robinson (1926: 237, 238) 
incorrectly gave the Nuttall number 564 for this 
collection. Taylor (1946: 116) repeated 
Robinson's error for the Nuttall number. 

N566 R270 

1 9 Amblyomma ovale 

ex. Speothas venaticus 

Charapaya (17'22'S, 66'45'W), Bolivia 

Publication: Robinson (1926: 29) gave the date as 

1900 and indicated that the collection contained 

only males. 

N567 R282 

Amblyomma albolimbatum [sex/stage not given] 

(2 d 1 , 1 9 Amblyomma albolimbatum) 
ex. Morelia variegata (Morelia spilotes variegatd) 
Perth (3 1 56'S, 1 1 550'E), W. Australia. 
L. E. Robinson det. 1926 

N568 R231 

Amblyomma [species epithet integrum crossed 

out and dissimile inserted]: [sex/stage not given] 

(7 d, 2 9, 7 N Amblyomma dissimile) 
ex. large snake 
Probably Frontera (18'32'N, 9238'W), Tabasco, 

May 1 . 1 .V. 1 90 1 [The full date is on vial label but 

not in catalogue] 

Publication: Robinson (1926: 167) indicated the 
date for this collection was V.I 900 and that only 
one female was present. 

N569 R267 

Amblyomma moreliae [sex/stage not given] [This 
species epithet crossed out and albolimbatum 
inserted] (1 9 Amblyomma albolimbatum) 


ex. Diemenia superciliosa (Diemenia textilis) 
Herdman's Lake, near Perth (31'56'S, 1 15'15'E), 

W. Australia 

Det. Neumann & Warburton 
Publication: Robinson (1926: 226) figured this 9; 
on p. 227 he incorrectly gave the collection 
number as N 1569. 

N570 R182 
2 cf Amblyomma marmoreum 
leopard tortoise 
No other data 
L. E. Robinson det. 

Note from Mr. Robinson 17. XI. 1922: 'They 
originally showed the characteristic ornamen- 
tation, sufficient of which is still visible, especially 
if examined under alcohol, to leave no doubt. The 
coarse punctations are larger than those seen in 
specimens off Rhinoceros in East Africa, and the 
dark ornamentation is more salient, but I do not 
consider these differences to be of specific impor- 
tance. The other morphological characters are 

N572 R288 

Haemaphysalis hystricis [sex/stage not given] (3 9 

Haemaphysalis hystricis) 
ex. Canis familiaris 
Lhassia, Assam, India (coordinates for Assam 

1907.R. A. Lorrain 

Note: Nuttall sent 1 cf, 1 9 of this collection to Or 
L. O. Howard of the U.S. Department of 
Agriculture. These specimens are now deposited 
in the Rocky Mountain Laboratory collection 
(RML 109001). 

Publications: Nuttall & Warburton (1915: 425); 
Hoogstraal et al. (19650: 476) gave combined 
collection data from Nuttall 572 and 573. 

N573 R293 

Haemaphysalis hystricis [sex/stage not given] (2 9 

Haemaphysalis hystricis) 
ex. Canis familiaris 
Lhassia, Assam, India (coordinates for Assam 

26WN, 93WE) 
1907.R. A. Lorrain 

Publication: Hoogstraal et al. (19650: 476) gave 
combined data from Nuttall 572 and 573. 

N574 R294C 

Haemaphysalis bispinosa [sex/stage not given] 

(10 cf, 6 9, 1 N Haemaphysalis bispinosa) 
No other data 

Haemaphysalis bispinosa is a vector of Kyasanur 
forest disease virus. 

N575 R243 

9 Haemaphysalis hystricis (2 9 Haemaphysalis 

Note: Nuttall 565 and 3330 also bear Rothschild 
number R243. 

N575x R163 

Haemaphysalis leachft [sex/stage not given] (1 cf 

Haemaphysalis indica)' 
ex. Hyaena crocuta (Hyaena hyaena) 
Darie (locality not verified) 
(Exp. Erlanger Hilgert) 

Note: The Nuttall number for this collection is 
575x in his catalogue but 575a on the vial label. 
Nuttall 1395 also bears Rothschild number R 163. 

N576 R294a 

Haemaphysalis hystricis [sex/stage not given] (1 9, 

1 N Haemaphysalis hystricis) 
ex. Canis familiaris 

Lushai Hills (23'10'N, 9250'E), Assam, India 
1907.R. A. Lorrain 

Publications: Nuttall & Warburton (1915: 425) 
indicated that N576 was composed of 1 cf, 3 9. 
Hoogstraal et al. (1965a: 476) saw the 9 labeled 
N576ii. The nymph, in a separate vial bears the 
label N576i. 

N577 R294 

Haemaphysalis leachi [sex/stage not given] (1 cf, 1 

9 Haemaphysalis leachi) 
ex. Ictonyx capensis (Ictonyx striatus) 
Bulawayo (2009'S, 2835'E), Zimbabwe 

Notes: The genus Ictonyx is also known in the 
literature as Zorilla. I. Geoffrey, 1826. 
Nuttall 595 also bears Rothschild number R294. 
Publication: Nuttall & Warburton (1915:471). 

N578 R154 

Haemaphysalis leachi (?) (not long enough) 

[sex/stage not given] (2 cf, 1 9 Haemaphysalis 


Deelfontein (20'17'S, 3238'E), Zimbabwe 
1. IV. 1 902 C.J.B. Grant 

Publication: Nuttall & Warburton (1915: 472) 
indicated the genus as Suricata. However, the 
collection locality appears to be out of the known 
range of the meerkat. 

N579 R212 

Haemaphysalis campanulata [sex/stage not 

given] (2 cf, 1 9 Haemaphysalis campanulata) 
ex. Canis familiaris 
Yokohama (3526'N, 139 8 37'E), Japan 
24. IV. 1902 Allan Owston 

Note: Nuttall 609 also bears the Rothschild 
number R212 but the data do not correspond to 

Publication: Nuttall & Warburton (1915: 472) 
indicated the collection contained 2 cf, 2 9 from a 



N580 R160 

1 cf 1 9 Haemaphysalis punctata 

[no host data] 

Cambridge (52 1 3'N, 00'08'E), England 

A. E. Shipley (Nuttall det.) 

Haemaphysalis punctata is a vector of the 

viruses Bhanja, tickborne encephalitis, Crimean 

Congo hemorrhagic fever, and perhaps Tribec. 

N581 R183 

1 N Haemaphysalis spinulosal [The species 
epithet crossed out and sp. inserted] (Probably 
H. leachi var. indica) (1 N Haemaphysalis sp.) 
(Bad condition, very queer. C. Warburton) 

ex. flying squirrel 

Bunguran (possibly Bungura 2620'N, 84'05'E), 

N582 R284 

Haemaphysalis bispinosa [sex/stage not given] (2 

9 Haemaphysalis bispinosa) 
ex. Equus caballus 

Lushai Hills (2310'N, 9250'E), Assam, India 
26. VI. 1 907 R. A. Lorrain 
Publication: Nuttall & Warburton (1915:431). 

N583 R291 

Haemaphysalis bispinosa [sex/stage not given] ( 1 

cf, 1 9 Haemaphysalis bispinosa) 
ex. Talpa sp. 

Lushai Hills (2310'N, 9250'E), Assam, India 
1907R. A. Lorrain 
Publication: Nuttall & Warburton ( 1 9 1 5: 43 1 ). 

N584 R294b 

Rhipicephalus haemaphysaloides [sex/stage not 

given] (3 cf, 1 9 Rhipicephalus 

No data 

N585 R242 

Rhipicephalus appendiculatus [sex/stage not 

given] (2 cf Rhipicephalus muehlensi) 
No data 

N588 R255 

Boophilus sp. incert. [sex/stage not given] (10 9 
Boophilus microplus) 

No host data 

Entre-Rios, Argentina. Probably Provincia de 
Entra Rios (32WS, 59WW), Argentina. 
(There are also two small populated areas with 
the name Entra Rios at 2318'S, 64 1 1'W and 
2850'S, 6411'W, which could be the collec- 
tion locality.) 

IV. 1 897 Dr C. Berg [C. Warburton det. 1918] 

Note: This collection also contained 1 cf of the 

human sucking louse Pediculus humanus. 

Boophilus microplus is a vector of the viruses 

Wad Medani, Seletar, and Crimean Congo 

hemorrhagic fever. 

N589 R290 

Argas persicus [sex/stage not given] (9 cf, 6 9, 5 N 

Argas persicus)) 
No history 

N590 R292 

Argas persicus [sex/stage not given] (11 cf , 4 9, 11 

N Argas persicus) 
No history 

Note: An additional 1 cf , 1 9 of this collection were 
presented to J. de Meza of Zomba, Malawi, 

N591 R214 

1 6L Argas vespertilionis 

ex. bat (also parasites off mouse) 

No particulars 

N592 R195 

3L Argas vespertilionis (& Nycteribia) 

ex. Scot 'us wroughtoni (Scotophilus temmincki 

Helwak(1722'N, 73'44'E), Maharashtra, India. 

N593 R224 

Argas vespertilionis [sex/stage not given] (3 L 

Argas vespertilionis) 
ex. Vesperugo pipistrellus (Pipistrellus 


Puttenham (5 1 49'N, 0044'W), England 

N594 R240 

Ornithodoros talaje [sex/stage not given] (14 N 

Ornithodoros capensis group) 
ex. nestlings 
Arrows Sand? Puhekiox, Hawaii (locality not 

30.VI.1900H.W. Henshaw 

N597 R153 

Ixodes hexagonus [sex/stage not given] (8 N 1 L, 

Ixodes hexagonus) 
ex. Erinaceus europaeus 
Belgium. E. A. Boulenger 
Publication: Nuttall & Warburton (1911:183). 
Ixodes hexagonus is a vector of tickborne 
encephalitis virus. 

N598 R206 

Ixodes hexagonus [sex/stage not given] (1 9, 6 N 

Ixodes hexagonus) 
ex. otter (Lutra vulgaris) (Lutra lutra) 
No particulars 

Publication: Nuttall & Warburton (1911: 182) 
indicated that the locality was England. 

N599 R209 

Ixodes hexagonus [sex/stage not given] (3 9, 4 N 

Ixodes hexagonus) 
ex. Erinaceus europaeus 
Cambridge (52 1 3'N, 0008'E), England 
Publication: Nuttall & Warburton (1911:182). 



N600 R221 

Ixodes hexagonus [sex/stage not given] (3 9, 1 L 

Ixodes hexagonus) 
ex. polecat Mustela putorius 
Boxworth (52'16'N, 0002'W), England 
Publication: Nuttall & Warburton (1911:1 82). 

N601 R208 

Ixodes hexagonus [sex/stage not given] (1 9, 41 L 

Ixodes hexagonus) 
ex. otter (Lutra vulgaris) (Lutra lutrd) 
No other data 

Note: One larva (RML 16476) of this collection 
was donated to the Rocky Mountain Laboratory 
by P. A. Buxton, London School of Hygiene and 
Tropical Medicine, in 1940. (See also Nuttall 

Publication: Nuttall & Warburton (1911: 182) 
indicated that the locality was England. 

N602 R222 

1 9 Ixodes hexagonus 

ex. Mustela erminea 

Tring (5 1 '48'N, 0040'W), England 


Publication: Nuttall & Warburton (1911:1 82). 

N603 R179a 

Ixodes hexagonus [sex/stage not given] (2 9, 1 N 

Ixodes hexagonus} 
ex. Mustela erminea 
Wigginton (5 1 47'N, 00'38'W), England 
4. V.I 903 Barrett coll. 
Publication: Nuttall & Warburton (1911:1 82). 

N604 R143 

Ixodes hexagonus [sex/stage not given] (4 N 

Ixodes hexagonus) 
ex. Mustela vulgaris (Mustela nivalis) 
Tring (5 1 48'N, 0040'W), England 
Publication: Nuttall & Warburton (1911:1 82). 

N605 R164 

Ixodes hexagonus [sex/stage not given] (3 9 

Ixodes lividus) 

ex. Cotile riparia (Riparia riparid) 
Worsborough Bridge (5332'N, 01'28'W), 


6.VII.1906A. Whitakercoll. 
Publication: Arthur (1963: 92) 
Ixodes lividus is a vector of tickborne 
encephalitis virus. 

N606 R225 

d"s, 9's, & larvae Ixodes hexagonus (3 9, 6 N, 8 L 

Ixodes hexagonus) 
ex. polecat (Mustela putorius) 
Aberystwyth (5225'N, 0405'W), Wales 

Publication: Nuttall & Warburton (1911: 183) 
gave the date as 1902. 

N607 R201 

Ixodes hexagonus [sex/stage not given] (1 N, 7 L 

Ixodes hexagonus) 
ex. Mustela erminea 
Tring (5 1 48'N, 0040'W), England 

Publication: Nuttall & Warburton (1911: 182) 
indicated that the collection was composed only 

N608 R215 

Ixodes hexagonus [sex/stage not given] (4 N 

Ixodes hexagonus) 
ex. Erinaceus europaeus 
Durnberg(4813'N, 1 1'55'E), Federal Republic of 


Publication: Nuttall & Warburton (1911: 183) 
gave the spelling as Dernberg rather than the 
catalogue spelling of Dermberg. 

N609 R212 

1 N Ixodes hexagonus 

ex. polecat (Mustela putorius) 

Aberystwyth (5225'N, 0405'W), Wales 


Note: Nuttall 579 also bears the Rothschild 

number R2 1 2 but the data do not correspond to 


Publication: Nuttall & Warburton (1911: 183) 

indicated that more than one nymph was in this 


N610 R219 

N's, larvae Ixodes hexagonus (1 N, 15 L Ixodes 

ex. Mustela erminea 
Tring (5 1 48'N, 0040'W), England 
Publication: Nuttall & Warburton (1911:1 82). 

N611 R145 

1 N, 1 L Ixodes hexagonus 

ex. Mustela erminea 

Lyndhurst (50'52'N, 1 34'W), England 

20.III.1901G. Tate 

Publication: Nuttall & Warburton (1911: 182) 

indicated that multiple nymphs and larvae were 

in this collection. 

N612 R180 

9's, N's Ixodes hexagonus (1 9, 7 N Ixodes 

ex. Mustela erminea 
Inismore (54 1 8'N, 07'34'W), Ireland 
Publication: Nuttall & Warburton (1911:1 82). 



N613 R202 

9's, N's Ixodes hexagonus (5 N Ixodes hexagonus) 

ex. stoat (Mustela ermined) 

No other data 

N617 R177 

1 9 Ceratixodes putus (Ixodes uriae) 

ex. wild duck 

No other data 

Publication: Nuttall & Warburton (1911:261). 

Ixodes uriae is a vector of the viruses tickborne 

encephalitis, Tuleniy, Oceanside, Zaliv 

Terpenya, Sakhalin, Tillamook, Avalon, Clo- 

Mor, Taggert, Bauline, Cape Wrath, Nugget, 

Tindholmur, Mykines, Great Island, Okhotskiy, 

Yaquina Head, Poovoot, Paramushir, and Runde. 

N618 R220 

1 cf, 1 9, 2 N Ceratixodes putus (Ixodes uriae) 

British. E. G. Wheler 

No particulars 

N619 R147 

1 d 1 , 2 9 Ixodes tenuirostris (2 rf, 2 9 Ixodes 

ex. Hypudaeus glareolus (Clethrionomys 


Campfer (4628'N, 0948'E), Switzerland 
7.VII.1904 [full date on vial label]. K. Jordan. 
Publication: Nuttall & Warburton ( 1 9 1 1 : 248). 
Ixodes trianguliceps is a vector of tickborne 
encephalitis virus. 

N621 R204 

N's Ixodes tenuirostris (1 N Ixodes trianguliceps) 

ex. Sorex vulgaris (Sorex araneus) 

Bevendean, (505 1 'N, 0005'E), England 


Publication: Nuttall & Warburton (1911: 248), 

indicated that more than one nymph was in the 


N622 R166 

9 Ixodes tenuirostris (1 9 Ixodes trianguliceps) 

ex. Arvicola amphibius (Arvicola terrestris) 



Publication: Nuttall & Warburton (1911: 248) 

indicated the collection was from England. 

N623 R197 

2 9 Ixodes tenuirostris (Ixodes trianguliceps) 
ex. Arvicola agrestis (Microtus agrestis) 
Tring (5 1 '48'N, 0040'W), England 
Publication: Nuttall & Warburton (1911: 248). 

N624 R149 

1 9 Ixodes tenuirostris (Ixodes trianguliceps) 

ex. Arvicola arvalis (Microtus arvalis) 

Campfer (4628'N, 0948'E), Switzerland 

20. VII. 1904 K.Jordan 

Publication: Nuttall & Warburton (1911: 248). 

N625 R148 

3 9 Ixodes tenuirostris (Ixodes trianguliceps) 

ex. Arvicola arvalis (Microtus arvalis) 

Campfer (4628'N, 0948'E), Switzerland 

5. VII. 1904 K.Jordan 

Publication: Nuttall & Warburton ( 1 9 1 1 : 248). 

N626 R226 

d"s, 9's Ixodes scapularis (4 rf, 4 9 Ixodes 

scapular is) 

ex. Cariacus virginianus (Odocoileus virginianus) 
Craven Co. (35'07'N, 7703'W), North Carolina, 

13.XI.1907 k [det. C. Warburton 5. 11.1909. Found 

with N666]' 

Publication: Nuttall & Warburton (1911: 158) 
gave the date of collection as XI. 1897. The date 
1 3.XI. 1 897 is also on the vial label. 

N633 R169 

N's Amblyomma sp. (8 N Amblyomma sp.) 

ex. Macropus sp. 

No other data 

Note: Nuttall 644 also bears the Rothschild 

number R169 and gives the additional data: W. 

Australia 28.VII. 1900. 

N634 R159 

Ixodes percavatus (2 9, 2 N Ixodes rothschildi 


ex. Puffin Fratercula arctica. Wrongly labelled Ix. 
putus '(Pocock)' O n back of label; no parti- 
culars. (Sent to Neumann who confirmed 
the determination II. 1909.) 
No other data. 

Publications: Nuttall & Warburton (1911: 221) 
described the 9 and N of Ixodes percavatus 
rothschildi. Zumpt (1952: 17) raised this taxon to 
specific rank; Arthur (1953: 222-226) indepen- 
dently raised /. percavatus rothschildi to specific 
rank; Arthur ( 1 963 : 1 34) referred to 2 9, 3 N in the 
type lot. 

There was (IX. 1977) a note in the vial with the 
words '1 N Missing'. 

N637 R213 

Ixodes bicornis [sex/stage not given] ( 1 1 9 Ixodes 

ex. Felis concolor 
'[American]' No other data 
Publication: Nuttall & Warburton (1911: 187) 
indicated only one 9 in the collection. 

N638 R276 

1 cf, 1 9 Ixodes loricatus 

ex. Didelphia (Didelphis sp.) 

Sapucay. Foster coll. 

Note: Locality is probably Sapucai (25 1 9'S, 

5655'W), Paraguay, which is in the range of /. 

loricatus. [See also Nuttall 1 39 1]. 

Publications: Nuttall & Warburton (1911: 269); 

Cooley& Kohls (1945: 190). 



N639 R256 

Ixodes loricatus [sex/stage not given] (1 d Ixodes 

ex. 'Piquoti' 
No other data 

N640 R241 

2 rf, 2 9, 2 N Ixodes loricatus (2 cf, 2 9 Ixodes 


ex. A teles melanochir (A teles g. geoffroyi) 
Frontera (18'32'N, 9238'W), Tabasco, Mexico. 

Note following locality '(Compared with 

Neumann's types at Toulouse, 1910; agrees 

fully but Nn's type larger).' 
Publications: Nuttall & Warburton (1911: 269); 
Cooley& Kohls (1945: 190). 

N641 R161 

Ixodes loricatus [sex/stage not given] (2 rf, 1 L 

Ixodes loricatus) 

ex. Didelphys aurita (Didelphis marsupialis) 
Brazil 24.XI.1901 

Publications: Nuttall & Warburton (1911: 269); 
Cooley& Kohls (1945: 190). 

N642 R165 

1 9, 1 N, 6 L Ixodes vestitus (1 9, 5 N Ixodes 

ex. Diemenia superciliosa (Diemenia textilis) 
Herdman's Lake, near Perth (31'56'S, 1 15'15'E), 

W. Australia 
B. H. Woodward 

Note: Nuttall 628 also bears Rothschild numfjer 

Publications: Nuttall & Warburton (1911: 255) 
indicated larvae as did the catalogue entry. These 
were missing 7.X.1977. Taylor (1946: 59, 60) 
repeated Nuttall & Warburton's description and 
figures. Roberts ( 1 960: 45 1 , 452). 

N643 R265 

2 N Ixodes holocyclus 

Perth (3 1 56'S, 1 1 5 1 5'E), W. Australia 

B. H. Woodward coll. 

Publication: Nuttall & Warburton (1911:238). 

N645 R275 

3 9 Ixodes australiensis 

ex. tame dog (Canis familiar is) 

Cranbrook (34 1 8'S, 1 1 732'E), West Australia 

1. IV. 1 900 J.T. Young 

Publications: Nuttall & Warburton (1911: 252). 

Taylor (1946: 57, 58) repeated Nuttall & 

Warburton's descriptions and figures. 

Note: This collection was seen by R. A. Cooley in 

1946 who made 3 slides of dissections (9 

spiracular plate; 9 legs I & IV; 9 hypostome) all in 

the Nuttall collection at the British Museum 

(N.H.). He gave them the Rocky Mountain 

Laboratory (RML) number 22284. 

N646 R210 

Ixodes australiensis [sex/stage not given] (8 9 

Ixodes australiensis) 
ex. Bettongia lescuri (Bettongia lesueuri) 
Kojonup (3350'S, 1 1 709'E), W. Australia 

Publications: Nuttall & Warburton (1911: 252). 
Taylor (1946: 57, 58) repeated Nuttall & 
Warburton's description and figures. Roberts 
(1960: 440); Roberts (1970: 40) gave the correct 
spelling of the locality. 

N647 R248 

3 9 Ixodes loricatus var. spinosus n. var. Nuttall, 

1 9 1 (2 9 Ixodes luciae) 
ex. large opossum (Didelphis sp.) 
Frontera ( 1 832'N, 92'38'W), Tabasco, Mexico 
May. Found with 648. Note long spine on coxa I. 
Note: These are the TYPE SPECIMENS of /. 
loricatus spinosus. 

Publications: Nuttall (1910: 411), original de- 
scription. Nuttall & Warburton (1911: 269), 
repeated original description and figures. 

N648 R248 

N's & larvae Amblyomma Icajennense (16 N 

Amblyomma cajennense) 
ex. large opossum (Didelphis sp.) 
Frontera ( 1 832'N, 9238'W), Tabasco, Mexico 
Note: Found with Nuttall 647 

N649 R185 

1 N Ixodes vespertilionis. 

ex. Vespertilio tricolor (Myotis tricolor) 

Kei Road, W. Kingwilliamstown, S. Australia [S. 

Australia crossed out and S. Africa inserted]. 

Kei Road, King William's Town (32'51'S, 

2722'E), South Africa. 
Purchased of Capt. H. Trevelyan. 
Publications: Nuttall & Warburton (1911: 277). 
Taylor (1946: 41) considered /. vespertilionis 
'very doubtfully Australian' and 'original 
specimen is unfortunately lost'. Hoogstraal (1956: 
569, 570) considered this collection to be of South 
African origin. 

Ixodes vespertilionis is a vector of the viruses 
Issyk Kul and tickborne encephalitis. 

N650 R203 

1 9 Ixodes fecialis TYPE (HOLOTYPE) 

ex. Dasyurus geoffroyi 

Cranbrook (34 1 8'S, 1 17'32'E), W. Australia 


Publications: Warburton & Nuttall (1909: 58) 

original description. Nuttall & Warburton (1911: 

248) repeated the original description and figures 

as did Taylor (1946: 56). Roberts (1960: 445) 

incorrectly gave the collection date as March 



N652 R168 

2 9 Ixodes unicavatm(\ 9, 1 N Ixodes unicavatus) 

ex. shag (Phalacrocorax aristotelis) 

Plymouth (50'23'N, 0410'W), England 


Publications: Nuttall & Warburton (1911: 266) 

indicated that the collection consisted of only one 

nymph. Arthur ( 1 963: 1 26) repeated this error. 

N666 R226 

1 9 Dermacentor nigrolineatus (1 9 Dermacentor 


ex. Cariacus virginianus (Odocoileus virginianus) 
Craven Co. (35 8 07'N, 7703'W), North Carolina, 

U.S.A. Found with N626 q.v. for details. 
C. Warburton det. 1917 

Dermacentor albipictus is a vector of Colorado 
tick fever virus. 

N667 R250 

9's, N's Ixodes unicavatus (39, 10 N, 3 L Ixodes 

ex. Phalacrocorax graculus (Phalacrocorax 


Cromarty (5740'N, 0402'W), Scotland. 
10.111. 1898 W.Roy 

Publications: Nuttall & Warburton (1911: 266): 
Arthur (1963: 126). 

N669 R178 

Haemaphy sails humerosa [sex/stage not given] 

TYPES (2 d 1 , 1 9, 1 N Haemaphysalis humerosa 


ex. Parameles macrura (Isoodon macrourus) 
Barrow Island (2048'S, 11523'E), N. W. 


Note: An additional 1 cf of this collection was sent 
to Dr L. O. Howard, U.S. Dept. of Agriculture, 
Washington, D.C., U.S.A. 2 1 .V. 1 909. 
Publications: Warburton & Nuttall (1909: 60) 
original description. Nuttall & Warburton (1915: 
496, 497) and Taylor (1946: 82) repeated original 
description and figures. Roberts (1963: 39) dis- 
cussed the type series and the note inserted in the 
tube by Nuttall that one specimen was missing 

N670 R172 

Haemaphysalis elongata [sex/stage not given] (1 d 1 
Haemaphysalis elongata, 4 d 1 , 2 9 Haema- 
physalis tiptoni) 

ex. Centetes ecaudatus (Tenerec ecaudatus) 


Note: The male of H. elongata has been given a 

collection number by the Rocky Mountain 

Laboratory (RML 71004) and by Harry 


Publications: Nuttall & Warburton (1915: 500). 

Hoogstraal (1953: 59, 64) discussed the two 

species represented in Nuttall & Warburton's 

(1915) figures and descriptions. 

N826 R189 

2 9 Ixodes hexagonus (Ixodes rubidus) 

ex. Coypu, Myopotamus coypus (Myocaster 


S. America. No further particulars 
Publication: Nuttall & Warburton (1911:1 83). ' 

N892 R295 

Haemaphysalis bispinosa [sex/stage not given] (7 

d, 7 9 Haemaphysalis bispinosa) 
ex. Canis familiaris 
Luthaia Mts, Assam 

Note: The locality is questionable. There are no 
Luthaia Mts as written in the catalog or Southaia 
Mts as in Nuttall & Warburton (1915: 431) to be 
found in gazetteers of India. I believe the locality 
to be Lushai Hills (2310'N, 9250'E), Assam, 
India as in N576, (R294a) and N582 (R284). 
Publication: Nuttall & Warburton ( 1 9 1 5: 43 1 ). 

N1066 R245 

5 9 Ixodes texanus var. (Ixodes sp. not /. 
texanus or /. hearlei) 

No other data 

N1067 R200 

1 9 Ixodes hexagonus 

ex. fox ( Vulpes vulpes) 

Boxworth (52 1 6'N, 0002'W), England 


Publication: Nuttall & Warburton (1911:1 82). 

N1068 R190 

6 d, 6 9 Ixodes canisuga (4 d 1 , 4 9 Ixodes lividus) 
Lyndhurst (5052'N, 0134'W), England 

11. VII. 1900 

Publications: Nuttall & Warburton (1911: 214). 
Arthur (1 963: 92) indicated 6 d, 6 9 present. There 
were 4 d, 4 9 in the collection as of IX. 1977. 

N1069 R146 

1 d, 2 9, 3 N Ixodes tenuirostris (Ixodes 


ex. Arvicola arvalis (Microtus arvalis) 
Campfer (4628'N, 0948'E), Switzerland 
1 1. VII. 1 904 Dr K.Jordan 
Publication: Nuttall & Warburton (1911: 248). 

N1105 None 

3 d 1 , 3 9, 6 N Dermacentor albipictus 

Blucher Hall (51'06'N, 12001'W), British 

Columbia, Canada 
III. 1910. Rec'd 2.IV.1910 from Hon. N. C. 

Rothschild (London). 

N1151 R296 

10 L Argas vespertilionis 

ex. P ipistrellus pipistrellus 

Oristano (39'54'N, 0835'E), Sardinia 

May 1907 DrA. Krausse 



N1152 R175 

6 LArgas vespertilionis 

ex. Vesperugo pipistrellus (Pipistrellus 


Tring (5 1 48'N, 0040'W), England 
1. IX. 1898 

Nil 53 R244 

2 L Argas vespertilionis 

No other data 

Ml 154 R271 

2 L Argas vespertilionis (Argas (Carios) sp.) 
Perth (3 1 56'S, 1 1 550'E), W. Australia 
Note: These specimens may represent a new 
species. It is not Argas australiensis, Argas daviesi 
or Argas dewae, the currently known representa- 
tives of the subgenus Carios in Australia. 

N1159 R142 

N's Ixodes hexagonus (3 N Ixodes hexagonus) 

ex. Mustela erminea 

Tring (5 1 48'N, 0040'W), England 


Note: One nymph (RML 16476) of this collection 

was donated to the Rocky Mountain Laboratory 

collection by P. A. Buxton, London School of 

Hygiene and Tropical Medicine, in 1940. (See 

also Nuttall 601). 

Publication: Nuttall & Warburton (1911:1 82). 

N1160 R194 

L's Ixodes hexagonus (17 L Ixodes hexagonus) 
ex. fox Vulpes vulpes (cub) 
Boxworth (52 16'N,0002'W), England 

Note: 2 additional larvae from this collection 
were sent to Prof. R. Matheson, Cornell Univer- 
sity, Ithaca, N.Y., U.S.A. , 27. V. 1929. 
Publication: Nuttall & Warburton (1911:182). 

N1161 R278 

9 Ixodes tenuirostris (Ixodes trianguliceps) 

Tarasp(4638'N, 10'25'E), Switzerland 

summer 1901 

Publication: Nuttall & Warburton (1911: 248) 

N1162 R207 

2 L Ixodes sp. 

ex. Dromicia concinna (Cercartetus concinnus) 

Note: The nocturnal marsupial C. concinnus 

inhabits the southern part of western and central 


N1163 R247 

L's Argas vespertilionis (1 3 L Argas vespertilionis) 

ex. Vesperugo pipistrellus (Pipistrellus 


Yalding (5 1 1 4'N, 0026'E), England. 
8. VIII. 1 897 [However, 1 997 was written] 
W. R. Ogilvie-Grant 

N1164 R250,251 

L's Argas sp. (5 L Argas boueti) 

ex. either Phalacrocorax graculus, Cromarty, or 

Megaderma cor, Salamana, Havish, 4,500 ft 

(Number on cork defaced) 
ex. Megaderma cor. 

Salamana, Havish 1,372 m (locality not verified) 
Notes: Megaderma cor ranges in East Africa from 
Ethiopia to Tanzania. 

The correct Rothschild number for this collection 
is R25 1 . For R250 see Nuttall 667. 

N1166 R230 

10 nymphs [genus & species not given] 


ex. Mus escularis (name not verified) 
St Aignan. Possibly Misima Island (10'41'S, 

15242'E), Louisade Archipelago. This island 

was formerly known as St Aignan. 
VIII-XI. 1897 (Meek) 

N1167 R186 


2 N Ixodes sp. [1 167], 1 d Hyalomma aegyptium 

[1 167a] (Hyalomma truncatum) 
ex. a 'Sassaby' a bastard hartebeest 
(Damaliscus lunatus) Republic of South Africa. 


Hyalomma truncatum is a vector of the viruses 
Dugbe, Bhanja, Jos, and Crimean Congo 
hemorrhagic fever. 

N1211 R237 

2 N Ixodes sp. (2 N, 1 L Ixodes fecialis) 

ex. Tarsipes rostratus or Smiothopsis murinus 

[vial label reads S. murinus] (Sminthopsis 


Albany (35'02'S, 1 1 753'E), W. Australia 
2 1. XII. 1 900 B. H. Woodward 

N1212 R272 

5 N Ixodes sp. (Ixodes holocyclus) 

Perth (3 1 56'S, 1 1 550'E), W. Australia 

B. H. Woodward 

No further details 

N1213 R216 

1 9 Ixodes fecialis var. aegrifossus (Ixodes fecialis) 

ex. Perameles obesula (Isoodon obesulus) 

Bannister (3240'S, 1 16'33'E), W. Australia 


Publication: Nuttall & Warburton (191 1: 250). 

N1386 R249 

N's, L's Amblvomma sp. (14 N, 2 L Amblyomma 

ex. Cariacus virginianus mexicanus (Odocoileus 


San Rafael (20'12'N, 965 1 'W), Veracruz, Mexico 
27. III. 1903 



N1387 R264 

1 N ? genus (Dermacentor andersoni) 

ex. Lagomys princeps (Ochotona princeps) 
Canadian National Park, ? Ottawa, Canada 
5. VIII. 1 899 J. F. Dippie 

Note: Ottawa, Ontario is far to the east of the 
ranges of D. andersoni and the pika. The 
Canadian National Park and the date 1899 
suggests a relationship to N1390 (R280), Banff 
National Park, which is within the range of both 
O. princeps and D. andersoni. 
Dermacentor andersoni is a vector of the viruses 
Powassan and Colorado tick fever, the bacterium 
of tularemia, and the rickettsia of Rocky 
Mountain spotted fever. 

N1389 R155 

larvae Ixodes sp. (3 N Ixodes sp.) 

ex. Crysomys longicaudatus (Oryzomys 


Valparaiso (3302'S, 7 1 38'W), Chile 
Date?J. A.Wolffshon 

N1390 R280 

2 N ? genus '(has eyes!)' (2 N Dermacentor 

Banff (5 1 38'N, 1 1 622'W), Alberta, Canada 

N1391 R269 

4 N ? genus (3 N Amblyomma sp.) 

ex. Didelphys sp. (Didelphis sp.) 

Sapucai (25'19'S, 5655'W), Paraguay 


N1392 R187 

N's Amblyomma sp. (10 N Amblyomma sp.) 
ex. Hare wallaby (Lagorchestes conspicillatus) 
Barrow Island (30'48'S, 115'23'E), N. W. 

N1393 R176 

larvae Amblyomma sp. (64 L Amblyomma sp.) 

ex. Bettongia penicillata 

Bokerup, W. Australia. Possibly Bokarup Swamp 

(34'24'S, 11650'E). 
20. IV. 1900 

N1394 R227 

2 N Amblyomma sp. 

ex. Ontalis vetula macalii (Neotoma alleni vetula) 

Frontera (18'32'N, 92'38'W), Tabasco, 

Switzerland [sic.] Mexico 

N1395 R163 

N's Rhipicephalus sp. (2 N Rhipicephalus sp.) 

ex. Hyaena crocuta (Hyaena hyaena) 

Darie (locality not verified) 

? Date. Von Erlanger 

N1396 R246 
1 N Amblyomma sp. 

ex. Black squirrel (Sciurus sp.) 

Frontera ( 1 832'N, 9238'W), Tabasco, Mexico 


N1397 R171 

larvae Amblyomma sp. (26 L Amblyomma sp.) 

ex. Macropus eugenii (Macropusfuligenosus) 

Cranbrook (34 1 8'S, 1 1 732'E), W. Australia 


N1398 R150 

1 larvae? genus (1 LHyalommasp.) 

ex. Gerbillus gerbillus 


1 .III. 1903 Hon. N. C. Rothschild 

""Locality not verified. However, N. C. Rothschild 
was in the Nile delta early in 1903. Quite 
possibly the locality is Zagazig (3035'N, 
31*3 l'E), Egypt. 

N1399 R162b 

N's Ixodes sp. (4 N, 4 L Ixodes texanus) 

ex. Putorius energermanis (Mustela vison) 

Sumas, B.C., Canada* 

III. 1903 

*Either Sumas Lake (49'07'N, 12202'W), British 

Columbia, Canada or Sumas (4900'N, 

122 8 13'W), Washington, U.S.A. 

N1497 None 

1 N Dermacentor sp. (Dermacentor albipictus) 

ex. Moose (Alces alces) 

New Brunswick, (47WN, 66WW), Canada 

30.XI.1911 J. Birrellcoll. 

N1498 None 

1 cf Dermacentor albipictus 

ex. man (Homo sapiens) (Dr Walker) 

New Brunswick (47WN, 66WW), Canada 

29.XI.1911 J. Birrellcoll. 

N1499 None 

2 c? Dermacentor albipictus 
ex. Moose (Alces alces) 

New Brunswick (47WN, 66WW), Canada 
30.XI.1911 J. Birrellcoll. 

N1500 None 

2 cf, 1 9, 1 N Dermacentor albipictus 

ex. carabou [sic.] (Rangifer tarandus) 

New Brunswick (47WN, 66WW), Canada 

25.XI.1911 J. Birrellcoll. 

N1501 None 

1 N Dermacentor sp. (Dermacentor albipictus) 

ex. carabou [sic.] (Rangifer tarandus) 

New Brunswick (47WN, 66'00'W), Canada 

25. XI. 1911 J. Birrell coll. 

N1502 None 

1 N Dermacentor sp. (Dermacentor albipictus) 

ex. carabou [sic.] (Rangifer tarandus) 

New Brunswick (47'00'N, 66WW), Canada 

25. XI. 19 11 J. Birrellcoll. 



N1503 None 

1 d Dermacentor sp. (5 N Dermacentor albipictus) 

ex. carabou [sic.] (Rangifer tarandus) 

New Brunswick (47'00'N, 66WW), Canada 

25.XI.1911 J. Birrellcoll. 

N1504 None 

1 9 Ixodes hexagonus (Ixodes cookei) '[Not 

typical bad condition; possibly /. rubidus]' 
ex. black cat 

New Brunswick (47WN, 66WW), Canada 
19.XII.1911 J. Birrellcoll. 
Ixodes cookei is a vector of Powassan virus. 

N3138 R926 

1 d Aponomma laeve var. capensis (Aponomma 

ex. Echidna sp. 
No further data 
Gift of Hon. N. C. Rothschild "(unmounted by 

me from a slide R926)". L. E. Robinson del. 
Note: This is not Aponomma concolor or A. 
oudemansi. Coxal spurring is similar to A. pattoni 
but the specimen lacks cervical pits. It is a species 
near A. pattoni. 

N3160 R238 

2 d 1 , 1 9, 1 N Dermacentor reticulatus ( 1 d 1 , 1 9, 1 
N Dermacentor reticulatus) 

Revelstoke, and Revelstoke Point, now known as 

Stoke Point (50 1 8'N, 040 1 'W), England 

Dermacentor reticulatus is a vector of the viruses 
tickborne encephalitis, Russian spring summer 
encephalitis, and Omsk hemorrhagic fever. 

N3330 R243 

1 9 Dermacentor auratus (Dermacentor sp.) 

No other data 

Note: Nuttall 565 and 575 also bear Rothschild 

number R243. 

N3502 None 

d's, 9's Dermacentor andersoni (7 d, 9 9 

Dermacentor andersoni) 
Okanagan Falls (49'21'N, 119'31'W), British 

Columbia, Canada 
V. 19 13 Rothschild coll. 
Gift of Seymour Hadwen, 1922 

N3537 None 

N's Haemaphysalis leachi (ION Haemaphysalis 


ex. Dipodillus campestris (Gerbillus campestris) 
Timgad (3530'N, 0635'E), Algeria 
1 5. IV. 1920 N. C. Rothschild & K. Jordan, coll. 
Publication: Hoogstraal (1955:222, 223) (see note 
under 3539). 

N3538 None 

N's Haemaphysalis leachi (UN Haemaphysalis 


ex. Dipodillus campestris (Gerbillus campestris) 
Timgad (35'30'N, 06 8 35'E), Algeria 
15.IV. 1920 N. C. Rothschild & K. Jordan coll. 

N3539 None 

N's Haemaphysalis leachi (4 N Haemaphysalis 


ex. Dipodillus campestris (Gerbillus campestris) 
Timgad (3530'N, 0635'E), Algeria 
1 7.IV. 1 920 N. C. Rothschild & K. Jordan coll. 
Publication: Hoogstraal (1955: 223) gave the 
Nuttall number 3739 for this collection. He 
indicated the total number of nymphs as 1 7 in 
combined collections 3537 and 3539. In 1977 the 
total number of nymphs in these two collections 
was 14. 

A single tick collection given by the Hon. 
Walter Rothschild to G. H. F. Nuttall via Guy 
A. K. Marshall 

N888 None 

2 d, 1 9 Hyalomma aegyptium (Hyalomma 


ex. giraffe (Giraffa camelopardalis subsp.) 
Ogo, Senegal 
ex. Riggenbach coll. given by Hon. Walter 

Rothschild to African Entomological 

Committee (rec'd 19.X.1909 from Guy A. K. 


Notes: There are two populated places and one 
campsite with the name Ogo in Senegal: PPL. 
1435'N, 1550'W; PPL. 1533'N, 1317'W; 
CMP. 1445'N, 1503'W. 

18 specimens of this collection were received 
11. X.I 909 by the Entomological Research 
Committee (Anon. 1910). 

Rothschild specimens missing from Nuttall collection 

N75 R1015, 1017, 1019 

3 d Amblvomma clvpeolatum ( = atrogenatum N. 

ex. tortoise 
Zoological Society's Gardens, London, 

11. X. 1895 

Presented by N. C. Rothschild 
Mounted (i) opaque & (ii) (iii) transparent on 3 

slides nos. 1015,1017,1019 
L. E.Robinson det. 26.111. 19 19 



N76 R964 
1 $ Amblyomma sp. 
ex Monitor 

Kalao Island. No further data 
Mounted opaque on slide No. 964 from N. C. 

N77 R1033, 1034, 1035 

3 cf Amblyomma marmoreum 

ex. Testudo pardalis (leopard tortoise) 

No further data. Mounted as transparent 

specimens on slides Nos. 1033, 1034, 1035 
Presented by N. C. Rothschild 
L. E. Robinson det. 26. III. 1919 

N78 R996,997,999 
1 cf, 2 9 Amblyomma albolimbatum 
ex. black & white snake 

30.1.1896. No further data. Mounted as trans- 
parent specimens Nos. 997, 996, 999 
Presented by N. C. Rothschild 
L. E. Robinson det. 26.111.1919. 

N90 R938-941 

4 d 1 Aponomma gervaisi var. lucasi 

ex. Ophiophagus bungarus (Hamadryad) 

Presented by N. C. Rothschild 

No particulars. Mounted on slides 938-941; 2 in 

glycerine, 2 in balsam 
C. Warburton det. 1.1915 

N91 R900 

1 cf Aponomma gervaisi var. lucasi 
ex. Varanus civitatus (Monitor) 
Presented by N. C. Rothschild 

No particulars. Mounted as slide 900 in dil. 
glycerin. C. Warburton det. 1. 1 5 

N92 R962,965 

2 c? Aponomma gervaisi 

ex. Varanus civitatus (Monitor) 
Presented by N. C. Rothschild 
No particulars. Mounted as slides 962, 965 in 
glycerin. C. Warburton det. 1.1915 

N93 R959,963,964,966 

4 d 1 , 1 9 Amblyomma decoratum 

ex. Monitor 

Kalas Island 

Presented by N. C. Rothschild 

No particulars. 5 slides, 3 in glycerin 2 cf in 

balsam: R959, 961, 963, 964, 966. C. 

Warburton det. 1.1915 

N216 R96 1-973 

9's, N's, L's Ixodes hexagonus 

ex. Lulra vulgaris 

No doubt British 

N. C. Rothschild 7 slides 

N217 R930,931 
2 9 Ixodes hexagonus 
ex. fox (Canis vulpes) 

No doubt British 

Gift of N. C. Rothschild (2 slides R930-93 1 ) 

N298 R1000-1011 

6 9, 5 N, 1 L Ixodes hexagonus 

ex. stoat (Putorius ermined) 

No doubt British 

Gift of N. C. Rothschild 12 slides 

N322 R929 
1 9 Ixodes hexagonus 
ex. Mustela putorius (ferret) 
Probably British 

Gift of N. C. Rothschild. R929 mounted in 
glycerin in glass cell. G. H. F. Nuttall det. 

N324 R944-946 
3 N Ixodes hexagonus 
ex. Erinaceus europaeus 
Probably British 

Gift of N. C. Rothschild (3 slides R944-946). G. 
H. F. Nuttall det. 

N395 R988-990 
3 L Ixodes unicavatus slightly fed 
ex. Phalacrocorax graculus (shag) 
No locality etc. Probably British 
Gift of N. C. Rothschild. Mounted on 3 slides 
R988-90. G. H. F. Nuttall det. 13.11.1915 

N396 R1032 

1 N Ixodes brunneusl 

ex. Erithacus rubecula (robin) 

No data. Probably British 

Gift of N. C. Rothschild 

Mounted on a slide R 1032 

N397 R911 
1 cf Amblyomma hirtum 
ex. bird (in spirit) 
Galapagos Islds, Pacific 
Ocean (W. of Ecuador) 
Gift of N.C.Rothschild 
Mounted on a slide R9 1 1 

L. E. Robinson det. 

N509 R982, 984-987 

4 9, 2 N Ixodes ricinus 

ex. Cervus capreolus (roe deer) 

? British 

Gift of N.C.Rothschild 

Mounted on 5 slides R982, 984-987 

N516 R901-3,905 

5 N Ixodes putus 

ex. Diomedea salvinii (albatross) 

No data 

Gift of N. C. Rothschild 5 slides 


N517 R934 
1 9 Ixodes putus 
ex. Uria lachrymans (ringed guillemot) 



No data 

Gift of N. C. Rothschild 1 slide 

N518 R975-979 

5L Ixodes putus 

ex. Puffin 

No data. Probably British 

Gift of N. C. Rothschild 5 slides 

G. H. F. Nuttall del. 12.11.1915 

N539 R1036 

d Haemaphysalis leachi 1 slide 

ex. Erinaceus albiventris 

Gebel Auli, White Nile, Egypt 

9.V.1900S. &F. Witherby 

Received from N. C. Rothschild 1908 

Det.G.H.F. Nuttall 27.1. 1909 

N544 R253 

Amblyomma sp. not varium; bad condition 

ex. 3-toed sloth (Bradypus tridactylus) 


N548 R257 

Aponomma gervaisi 

ex. Boa 

Feb. 1887 

Given to D. S. MacLagan 14/10/38 

N551 R181 
Amblyomma triguttatum 
ex. Kangeroo(sic.) 
Barrow Island, N. W. Australia 
20.XI.1900C.J. T. Tanney 

N571 R279 

Aponomma sp. 

ex. Diemenia superciliosa 

Herdman's Lake, Perth, West Australia 

N586 R193 

d Rhipicephalus sanguineus 

ex. Camel 

Locality unknown 

N587 R151 

2 d Rhipicephalus sanguineus 

Zaghig, 25.11. 1908 

N.C.Rothschild coll. 

N595 R294 
Ornithodoros moubata 
No history 

Note: Nuttall 577 also bears Rothschild number 

N596 R157 
N's Ornithodoros megnini 
Okanagan, British Columbia 
1 2. XII. 1902 Allan Brooks 

Note: Otobius is the correct genus for this tick 

N620 R942 

1 9 Ixodes tenuirostris 

ex. Mus minutus (Harvest mouse) 

No further particulars '[Mounted on a slide]' 

N628 R165 

9 & N Ixodes ricinus 

No particulars 

Note: Nuttall 642 also bears Rothschild number 


N629 R233 
4 d Ixodes ricinus 
ex. stag 
No particulars 

N630 R217 

3 N Ixodes ricinus 
ex. Sciurus vulgaris 

17. IV. 1901 Timmer, Ireland 

N631 R223 

4 9 Ixodes ricinus 

ex. Erinaceus europaeus 
Belgium E. A. Boulengercoll. 

N632 R173 
2 9 Ixodes ricinus 
ex. fox 
No particulars 

N644 R169 

2 9 Ixodes holocyclus 

ex. Macropus sp. 

W.Australia 28. VII. 1900 

Note: Nuttall 633 also bears Rothschild number 


N656 R205 

2 L Ixodes ricinus 
ex. Mustela erminea 
Lyndhurst, Hants., England 
25. VII. 1 900 G. Tatecoll. 

N674 R922-924 

3 9 Ixodes hexagonus 
ex. Myopotamus coypu 

? locality, host a rodent from S. America 
Presented by N. C. Rothschild (R922-924) '[see 
record of 2 other 9'$ in Ticks Part II, p. 183]' 
G.H.F. 19 15 

N746 R232 
larvae Ixodes hexagonus 
ex. polecat 
Aberystwyth, Wales 
VIII. 1900 

N1388 R162 

1 LArgassp. 

ex. Putorius energermanis 

Sumas, B.C., Canada 


N3136 R958 
1 d Hyalomma sp. 

20 -J- E. KEIRANS 

ex. Capra falconer i (the Markhoor) 

No further data 

Gift of N. C. Rothschild (slide R958) 

Note: The host occurs in Kashmir, Afghanistan, 

Gilgit, etc. 

N3139 R1012-1020 

Hyalomma syriacum 

ex. tortoise 

No further data 

Gift of N. C. Rothschild (5 slides i-v 


N3140 R1022, 1023 

2 9 Hyalomma syriacum 

ex. Testudo pardalis 

No further data 

Gift from N. C. Rothschild 

R1022 1023 

(2 slides, 

N3141 R906, 1031 

Hyalomma syriacum 

ex. Araconda [sic] 

No further data (evidently bought slides from M. 
Pillischer, optician, 88 New Bond St., London 
W.) Gift of N. C. Rothschild (2 slides R906, 

N3157a R1026-1029 

N(3 1 57) 3 d Rhipicephalus evertsi 

(a) 1 cf Boophilus decoloratus 

ex. Zebra No locality 

Gift of N.C.Rothschild 1908 

Mounted on slides in balsam (R 1026-1 029) 

R. evertsi det. C. Warburton 18.111.1915 
B. decoloratus det. C. Warburton & G. 
Nuttall 1915 

H. F. 

N3158 R907 

1 9 Rhipichephalus haemaphysaloides 


Mounted on slide in glycerin 

Gift of N.C.Rothschild 1908 

(Prep, bought from Fredk. Enock*) 

*Spelled Enock in this collection, but see N3 1 59 

N3159 R908-910 

3 N Argas reflexus 


3 slides, glycerine in cells G. H. F. Nuttall det. 


Gift of N.C.Rothschild 1908 
(3 slides Nos. 908-9 10) Fredk. Enoch prep. 
Cost 2 shillings each! 

N3351 None 

9 Amblyomma varium 

ex. Bradypus tridactylus (sloth) 

Gift of N. C. Rothschild years ago. 

Scutum, legs, spiracle on 3 slides 

L. E. Robinson det. 26. III. 1919 

N3536 None 

1 9 Argas persicus 

Found on wall 

Djama, Algeria 


N. C. Rothschild & K. Jordan coll. 

Rothschild specimens donated to British Museum (Natural History) 

Species of Ixodoidea represented in the collection of the British Museum (Natural 
History), donated by the Hon. N. C. Rothschild 

K numbers follow each species. Numbers in parentheses refer to missing collections. Thus, 
validity of original determinations is unverified. 


Argas persicus (Oten)Kl, K2,(K148) 
Argas reflexus (Fabricius) K3 
Argas sanchez i Duges K.4 

Ornilhodoros talaje Guerin-Meneville K5 


Amblyomma albolimbatum Neumann K6 
Amblyomma americanum (Linnaeus) K7 
Amblyomma brasiliense Aragao K.8 

Amblyomma cajennense (Fabricius) K9 
Amblyomma calcaratum Neumann K10 
Amblyomma compressum Macalister Kl 1 
Amblyomma gemma Donitz K12, K13,(K149) 
Amblyomma humeraleC. L. Koch K14, K15 
Amblyomma incisum Neumann K16 
Amblyomma longirostre(C. L. Koch) K17, K18 
Amblvomma maculatum C. L. Koch K19, K20, 


Amblyomma naponense (Packard) K23 
Amblyomma nodosum Neumann K8, K24, K25 



Amblyomma ovale C. L. Koch K8, K23, K26, 

K27, K28, K29, K30, K3 1 , K32 
Amblyomma postoculatum Neumann K33 
Amblyomma pseudoconcolor Aragao K34 
Amblyomma testudinarium C. L. Koch K35 
Amblyomma variegatum (Fabricius) K36, K37 
Amblyomma sp. K38, K39 

Aponomma concolor Neumann K40, K41, K42, 

Aponomma decorosum (L. Koch) K44, K45, 

K46, K47 

Aponomma exornatum (C. L. Koch) K48 
Aponomma fimbriatum (C. L. Koch) K49 
Aponomma gervaisi var. trimaculatus (Lucas) 


Aponomma latum (C. L. Koch) K50 
Aponomma trimaculatum (Lucas) K5 1 

Boophilus decoloratus (C. L. Koch) K52 
Boophilus microplus (G. Canestrini) K53 
Dermacentor atrosignatus Neumann group K54 
Dermacentor reticulatus (Fabricius) K55, 

Haemaphysalis bancrofti Nuttall and Warburton 

K57, K58, K59, K60 
Haemaphysalis calcarata Neumann K6 1 
Haemaphysalis juxtakochi Cooley K53 
Haemaphysalis lagostrophi Roberts K33 
Haemaphysalis leachi (Audouin) K62, K63, K64, 

K66, K67, K68, K69, K70, K71, K72, K73, 

K74, K75,(K151) 

Haemaphysalis leachi indica Warburton (Kl 52} 
Haemaphysalis leachi (Audouin) group K76 
Haemaphysalis novaeguineae Hirst K77, K78, 


Haemaphysalis parmata Neumann K80 
Haemaphysalis spinigera Neumann K8 1 
Haemaphysalis spinulosa Neumann K82, K83 
Haemaphysalis sulcata G. Canestrini, and 

Fanzago K84, K85 

Haemaphysalis tibetensis Hoogstraal K86 

Hyalomma aegyptium (Linnaeus) K87 
Hyalomma anatolicum excavatum C. L. Koch 


Hyalomma marginatum C. L. Koch K89 
Hyalomma truncatum C. L. Koch K90, (Kl 53) 
Hyalomma sp. K9 1 , K92, K93 

Ixodes alluaudi Neumann K94, K95, K96, K97, 


Ixodes cordifer Neumann K99, K100, K101, 

Ixodes fecialis Warburton and Nuttall K103 

Ixodes hexagonus Leach K104, K105, K106, 

K107, K108,K109,K110 
Ixodes loricatus Neumann Kl 1 1 , Kl 12 
Ixodes luciae Senevet Kl 1 3 
Ixodes ornithorhynchi Lucas K 1 1 4 
Ixodes cumulatimpunctatus Schulze Kl 15 
Ixodes ricinus (Linnaeus) Kl 16, (Kl 54) 
Ixodes tasmani Neumann Kl 1 7 
Ixodes thomasae Arthur and Burrow K69 
Ixodes trianguliceps Birula Kl 18 
Ixodes vespertilionis C. L. Koch K119, K120, 

K121, K122, K123, K124,(K155) 

Rhipicephalus haemaphysaloides Supino K125 
Rhipicephalus jeanneli Neumann K126 
Rhipicephalus kochi Donitz K127 
Rhipicephalus kochi Donitz group K128 
Rhipicephalus longicoxatus Neumann K127 
Rhipicephalus pravus Donitz K127, K129, K130, 


Rhipicephalus pulchellus (Gerstacker) K93, K132 
Rhipicephalus sanguineus (Latreille) K133, 

K134, K135, K136, K137, K138, K139, K140, 


Rhipicephalus simusC. L. Koch K142, K143 
Rhipicephalus sp. K144, K145, K146, K147 

Rothschild specimens in British Museum (Natural History) collection with type 


B.M.(N.H.) Accession No. 

19 14. 1 1.1 7. 1-46. 


Dermacentor reticulatus aulicus Hirst, 1916. Ann. Mag. nat. Hist. 

(8), 17:308. 

Haemaphvsalis spinigera novaeguineae Hirst, 1914. Trans, zool. 

Soc. Lorn/ 20: 325, fig. 16. 

Haemaphvsalis tibetensis Hoogstraal, 1965. J. Parasit. 51: 452, 

figs. 1-26." 



Rothschild specimens in British Museum (Natural History) collection 


Kl 1912.6.21.21 

1 cf Argas persicus 

In railway carriage 

El Kantara (35 13'N,05'43'E), Algeria 


Hon. Walter Rothschild & Ernst Hartert 

Argas persicus is a vector of Crimean Congo 

hemorrhagic fever virus. 

K2 1919.8. 14.1-70 (pt) 

1 $ Argas persicus 

No host 

Ouargla house (locality not verified) 

E. Hartert and C. Hilgert 

Note: The locality is probably a house in Ouargla 

(3 r57'N,0520'E), Algeria. 

K3 1912.6.21,1-2 

1 9, 1 N Argas reflexus 

ex. Columba livia 

Leipzig (5118'N, 1220'E), German Democratic 

V.I 9 10 
O. Fritsche 

Argas reflexus is a vector of the viruses Ponteves 
and Grand Arbaud. 

K4 1912.12.4.1-8 

Argas persicus (1 cf , 5 9, 3 N Argas sanchezi) 

ex. chicken 

Price, Pinal Co. (3305'N, 1 1 1'30'W), Arizona, 

Dr R. E. Kunzr 

K5 None 

1 cf , 3 9, 3 N Ornithodoros talaje 

ex. native house 

Colombia, S. America 


K6 1912.6.21.304-318 

Amblyomma albolimbatum (19 cf, 6 9 
Amblyomma albolimbatum; 1 9 Haema- 
physalis sp.; 1 9 Haemaphysalis sp.) 

ex Trachysaurus rugosus 

Bernier Is. (2452'S, 1 1308'E), W. Australia 


B. H. Woodward 

Note: 1 cf, 1 9 included in the above total were 

seen by R. A. Cooley in 1946 who made six slides 

of dissections (RML 22306). 

The Haemaphysalis specimens represent either 

two species or a single exceptionally variable 

species (Hoogstraal, personal communication). 

K7 None 

Amblyomma sp. (1 N Amblyomma americanum) 

ex Scuirus niger 

Mount Pleasant (3247'N, 7952'W), South 

Carolina, U.S.A. 

A. T. Wayne 

Amblyomma americanum is a vector of Lone Star 

K8 None 

Amblyomma nodosum (6 cf, 1 9 Amblyomma 

brasiliense; 25 cf, 5 9 Amblyomma nodosum; 1 9 

Amblyomma ovale) 
Host unknown 
Joinvile (2618'S, 48'50'W), Humboldt, Estadode 

Santa Catharina, Brazil 
W. Ehrhardt 

K9 1912.6.21.326-337 

6 cf, 1 1 9, 3 N Amblyomma cajennense 

ex. cattle, Canis familiaris & Homo sapiens 

St. Ann's ( 1 82 1 'N, 77 1 6'W), Jamaica 

22-29. III.1911 

F. E. Sherlock 


Amblyomma concolor! '9 etc.' (79, 1 N, 4 L 

Amblyomma calcaratum) 
ex. Tapir americanus ( Tapirus terrestris) 
Joinvile (26 1 8'S, 4850'W), Humboldt, Estado de 

Santa Catarina, Brazil 
W. Ehrhardt 15. IX. 19 12 

Kll 1912.6.21.299-302 

Amblyomma cuneatum (Species epithet crossed 

out and compressum inserted) (1 cf, 4 9 

Amblyomma compressum) 
ex. Pangolin (or Scaly Ant-Eater) Manis sp. 
Bibianha, Gold Coast. Bebianiha (0543'N, 

0027'W) or other populated places of this 

name nearby. Ghana 
9. XII. 1911 
H. G. F. Spurrell 

Note: Vial label gives accession number as 

K12 1911. 12.9.1-320 pt 

2 9 Amblyomma gemma 

ex. Oxen 

Voi (0323'S, 2834'E), Kenya 

25. IV. 1910 

Robin Kemp 

K13 191 1.12.9. 1-320 pt 

5cf , 4 9 Amblyomma gemma 

ex. Oxen 

Voi (0323'S, 3834'E), Kenya 

30.IV. 1910 

Robin Kemp 



K14 1912.10.31.18-20 
1 cf, 3 9 Amblyomma humerale 
ex. donkey (Equus asinus) 
Locality unknown 


4 cf Amblyomma humerale 

ex. turtle 



1 cf Amblyomma incisum 

ex. tapir ( Tapirus sp.) 


Note: There is 1 9 in this collection which is 

tentatively being called A. incisum but which has 

a 3/3 hypostome. 

K17 1912.6.21.338 

1 cf Amblyomma longirostre 

ex. Galera barbara (Eira barbara) 

Mar de Espanha (21'52'S, 43'00'W), Minas 

Gerais, Brazil 
Z. F. Zikan 

K18 1912.6.21.339-343 

2 cf, 3 9 Amblyomma longirostre 

ex. Synetheres prehensilis (Coendou prehensilis) 
Mar de Espanha (21'52'S, 43WW), Minas 

Gerais, Brazil 
Z. F. Zikan 

K19 1912.10.31.13 

1 cf Amblyomma maculatum 

ex. Canis sp. 

Buenos Aires (3436'S, 5827'W), Argentina 


Miss Idina G. Runnael 

K20 1912. 12.4.12-14 

1 cf, 1 $ Amblyomma maculatum 

ex. Small Chubut Fox 

BahiaBlanca(3843'S, 62 17'W), Argentina 


E. Weiske 

K21 1919.8. 14.1-70 pt 

4 cf, 5 9 Amblyomma maculatum 

ex. Blastocerus dichotomus (B. bezoarcticus) 

Parana Delta (vicinity of 3343'S, 59'15'W), 

Robin Kemp 

K22 None 

3 cf Amblyomma maculatum 

ex. Pseudalopex gymnocerus (Dusicyon 

Laguna Alsina (3649'S, 62'13'W), to Buenos 

Aires (3436'S, 5827'W), Argentina 
Robin Kemp 

K23 1919.8. 14. 1-70 pt 

Amblyomma mantiquirense (1 cf Amblyomma 


ex. Hydrochaerus hydrochaeris 
Eastern Ecuador 

This collection also contains 1 9 of Amblyomma 
ovale or a closely related species. 

K24 None 

1 cf, 1 $ Amblyomma nodosum 

ex. Anteater 

Colombia, S. America 


K25 1912.6.21.325 

1 cf, 3 N, 3 L Amblyomma nodosum 

ex. Myrmecophaga tetradactla (Myrmecophaga 


Misantla (1956'N, 9650'W), Mexico 

W. Engelmann 
Note: Immature specimens tentatively this sp. 

K26 1912.6.21.344-352 

8 cf, 4 9, 3 N Amblyomma ovale 

ex. Nasua socialis (Nasua nasud) 

Mar de Espanha (2152'S, 43'00'W), Estado de 

Minas Gerais, Brazil 
Z. F. Zikan 

Note: Also present were 1 cf, 2 9 of the 
mallophagan Neotrichodectes pallidus 

K27 1912.6.21.353-358 

7 cf , 3 9 Amblyomma ovale 

ex. Galera barbara (Eira barbara) 

Mar de Espanha (20'52'W, 43WW), Minas 

Gerais, Brazil 
Z. F. Zikan 

K28 1921.12.9.10-11 

1 cf, 1 9 Amblyomma ovale 

ex. Galictes vittata (Grison vittatd) 


W. Ehrhardt 

K29 1921.12.9.12-20 

8 cf , 2 9 Amblyomma ovale 

ex. Nasua familiar is (Nasua nasua) 
Joinvile(2618'S,4850'W), Humboldt, Estado de 

Santa Catharina, Brazil 
W. Ehrhardt 

K30 None 

3 cf , 2 9 Amblyomma ovale 

ex. Procyon cancrivorus 

Joinvile (26'18'S, 48'50'W), Humboldt, Estado de 

Santa Catharina, Brazil 
W. Ehrhardt 



K3 1 None 

1 rf A mblyomma ovale 

ex. Nasua socialis (Nasua nasua) 

Mar de Espanha (21'52'S, 43WW), Estado de 

Minas Gerais, Brazil 
Z. F. Zikan 

K32 None 

1 2 rf, 5 9 Amblyomma ovale 

ex. Nasua socialis (Nasua nasua) 

Mar de Espanha (21'52'S, 43WW), Estado de 

Minas Gerais, Brazil 
Z. F. Zikan 

K33 1912.6.21.319-324 

Amblyomma albolimbatum (2 9 Amblyomma 

postoculatum; 5 9 Haemaphysalis lagostrophi) 
ex. Lagostrophus fasciatus 
Bernier Island (2452'S, 1 1 308'E), W. Australia 
B. H. Woodward 

K34 None 

Amblyomma concolor (2 rf, 7 N, 3 L Amblyomma 

ex. Armadillo 

La Rioja (2926'S, 66 1 5'W), Argentina 
E. Giacomelli 

K35 1912.12.4.15-16 

2 d Amblyomma testudinarium 

ex. Sheep 

Sri Lanka 

K36 None 

1 N Amblyomma variegatum 

ex. Raven 

Harar (09 1 9'N, 4209'E), Ethiopia 


G. Kristensen 

Amblyomma variegatum is a vector of the viruses 

Dugbe, Thogoto, Jos, Crimean Congo 

hemorrhagic fever, Nairobi sheep disease, Bhanja 

and Somone. 

K37 None 

1 d Amblyomma variegatum 

Host unknown 

Harar (09 1 9'N, 4209'E), Ethiopia 


G. Kristensen 

K38 None 

1 N Amblyomma sp. 
ex. Canis familiar is 

Harar (09 1 9'N, 4209'E), Ethiopia 


G. Kristensen 

K39 None 

2 N Amblyomma sp. 

ex. Homo sapiens 

Gola National Forest (0656'N, 1045'W), Liberia 


R. H. Bunting 

K40 1912.6.21.370-380 

Amblyomma concolor (3 9, 23 N Aponomma 


ex. Echidna aculeata (Tachyglossus aculeatus) 
Gippsland District (3800'S, 147WE), Australia 
A. Coles 

K41 None 

Amblyomma concolor (11 N Aponomma 


ex. Echidna sp. (Tachyglossus aculeatus) 
A. Coles 

K42 1912.6.21.381-383 

Amblyomma concolor (3 9, 3 N Aponomma 


ex. Echidna aculeata (Tachyglossus aculeatus) 
Traralgon (3812'S, 14632'E), Australia 
A. Coles 

K43 1912.6.21.384-386 

Amblyomma concolor (3 9 Aponomma concolor) 

ex. Echidna sp. (Tachyglossus aculeatus) 



A. Coles coll. 

K44 None 

Aponomma decorosum (Species epithet crossed 

out and undatum inserted) (22 cf, 3 9, 3 N 

Aponomma decorosum) 
Ex. Varanus sp. 
Kelsey Creek (2026'S, 14827'E), Queensland, 

A. Fontaine 

K45 1912.6.21.391^01 

Aponomma decorosum (Species epithet crossed 

out and undatum inserted) (22 cf , 6 9, 18 N, 2 L 

Aponomma decorosum) 
ex. Brown Iguana ( Varanus sp.) 
Kelsey Creek (2026'S, 14827'E), Queensland, 

A. Fontaine 
Publication: Kaufman (1972: 369) 

K46 19 12.6.2 1.402^10 

Aponomma decorosum (Species epithet crossed 

out and undatum inserted) ( 1 5 cT, 1 3 9, 2 1 N, 2 L 

Aponomma decorosum) 
ex. Black Iguana ( Varanus sp.) 
Kelsey Creek (2026'S, 14827'E), Queensland, 




A. Fontaine 

Publication: Kaufman (1972: 369) 

K47 None 

Aponomma decorosum (Species epithet crossed 

out and undatum inserted) (1 d 1 , 1 9, 4 N 

Aponomma decorosum) 
ex. Iguana ( Varanus sp.) 
Publication: Kaufman (1972: 370) 

K48 1912.10.31.21-25 

2 cf Aponomma exornatum 

ex. Egyptian monitor (Varanus sp.) 

Bibianha [Bebaianiha (05'43'N, 0027'W) or 

other populated place of this name nearby], 

H. G. F. Spurrell 

K49 1912.6.21.387-390 

Aponomma ecinctum (2 d 1 , 2 9 Aponomma 


ex. Tiger snake (Notechis scutatus) 
Melbourne (3749'S, 14458'E), Australia 
A. Coles 

Note: Very poor specimens; hypostomes lacking 
in all specimens. Capitulum present in only one 9 
but from porose areas and scutal configuration of 
9 and scutum and tarsi of cf these appear to be 
good A. fimbriatum. 

K50 1912.6.21.416 

1 d Aponomma latum 
ex. Hystrix sp. 

Mduna River, Hlabisa District, (2810'S, 
3210'E), Zululand Region, Republic of South 


F. Toppin 

Received from Natal Museum, Pietermaritzberg 

H. Hoogstraal det. 1952 'unusual host' 

K51 1912.6.21.359-369 

Aponomma gervaisi var. trimaculatus (11 d, 2 9, 

1 N Aponomma trimaculatum) 
ex. Python sp. 
Satte'lberg, Huon Gulf (0710'S, 147'25'E), 

Papua New Guinea 
Prof. F. Forster 

K52 None 

2 9 Boophilus decoloratus 
Host unknown 

Dr E. J. Baxter 

Boophilus decoloratus is a vector of the viruses 
Dugbe, Thogoto, Bhanja, Jos, Somone and 
Crimean Congo hemorrhagic fever. 

K53 1912.10.31.26-29 

Haemaphysalis kochi(\ 9 Boophilus microplus; 2 

d, 1 9, 1 N Haemaphysalis juxtakochi) 
ex. Deer 
Caparo (10'27'N, 6120'W), Trinidad Island, 

Trinidad and Tobago 
S. M. Klages 

K54 1912.12.4.17 

1 cf Dermacentor atrosignatus group 
ex. Sheep 

Sri Lanka 

K55 None 

Dermacentor reticulatus var. aulicus SYNTYPE 

( 1 9 Dermacentor reticulatus) 
Host unknown 
Valescure, nr. St. Raphael (4326'N, 0646'E), 


Publication: Hirst (1916: 308) original 

K56 1912.6.21.77-97 

2 d, 8 9 Dermacentor reticulatus 
ex. Canis familiaris 

Kuybyshev (5540'N, 7812'E), Tomsk, U.S.S.R. 


W. M. Thomas 

K57 1912.6.21.69-71 

1 9 Haemaphysalis bancrofti 

ex. Horse Equus caballus 

Kelsey Creek (2026'S, 14827'E), Queensland, 

A. Fontaine 

K58 1912.6.21.76 

1 d Haemaphysalis bancrofti 

ex. Black Wallaby [Macropus dorsalis, the black- 
striped wallaby or Wallabia bicolor, the black, 
or black-tailed wallaby, both of which occur in 

Kelsey Creek (2026'S, 14827'E), Queensland, 


A. Fontaine 

K59 None 

2 d Haemaphysalis bancrofti 

ex. Wallaby ( Wallabia sp. or Macropus sp.) 
Kelsey Creek (2026'S, 14827'E), Queensland, 

A. Fontaine 

K60 None 

2 d Haemaphysalis bancrofti 
ex. Canis familiaris 



Kelsey Creek (20'26'S, 14827'E), Queensland, 

A. Fontaine 

K61 None 

2 rf Haemaphysalis calcarata 

Host unknown 

Voi (0323'S, 3834'E), Kenya 


Robin Kemp 

K62 None 

Haemaphysalis leachi leachi (2 d 1 , 2 9 Haema- 
physalis leachi) 
Host unknown 
Dr E. J. Baxter 

K63 1912.10.31.30-35 

Haemaphysalis leachi leachi (9 d 1 , 2 9 Haema- 
physalis leachi) 
ex. Felis sp. 

Lake Suai, Ethiopia (locality not verified) 
[Hirst original det. Haemaphysalis sp.] 

K64 1912.6.21.33-35 

2 rf, 2 9 Haemaphysalis leachi 

ex. Hyaena (Hyaena hyaena) 

Harar (09 1 9'N, 4209'E), Ethiopia 


G. Kristensen 

K65 1912.6.21.36^4 

1 d 1 , 8 9 Haemaphysalis leachi 

ex. Felis catus 

Harar (09 1 9'N, 4209'E), Ethiopia 


G. Kristensen 

K66 19 12.6.2 1.4 5-49 

5 cf Haemaphysalis leachi 

ex. Marten 

Harar (09 1 9'N, 42'09'E), Ethiopia 


G. Kristensen 

K67 1912.6.21.50-55 

5 cf, 3 9 Haemaphysalis leachi 

ex. 'Badlington' dog 

Gola National Forest (06'56'N, 1045'W), Liberia 


R. H. Bunting 

K68 1912.6.21.56-58 

1 d 1 , 2 9 Haemaphysalis leachi 

ex. Felis serval 

Mpwapwa (062 1 'S, 3629'E), Tanzania 

15.X. 1911 

Marschner. Rec'd. from W. Fasius 

K69 None 

Haemaphysalis leachi (4 d Haemaphysalis leachi; 

2 9 Ixodes thomasae) 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 

K70 None 

7 rf, 109 Haemaphysalis leachi 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 

K71 None 

4 rf, 4 9 Haemaphysalis leachi 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 

K72 None 

2 cf Haemaphysalis leachi 

ex. Domestic Rabbit (Oryctolagus sp. probably.) 

The genus Oryctolagus does not exist south of 

the Sahara unless bred in captivity. 
Rombo, Kilimanjaro (0305'S, 3720'E), 

20. VI. 1910 
Robin Kemp 

K73 None 

1 1 d 1 , 8 9, 1 N Haemaphysalis leachi 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 

K74 None 

25 d 1 , 1 1 9 Haemaphysalis leachi 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 

K75 None 

1 cf Haemaphysalis leachi 

ex. Genetta stuhlmanni 

Fennek, Mutoragwa 

Aberdare Mts, Kenya (locality not verified*) 

23. HI. 19 10 

Robin Kemp 

*There is a Mutarakwa Hill (0047'S, 3514'E) in 

the Aberdares that is a possible locality for this 




K76 1912.6.21.59-65 

Haemaphysalis leachi! (7 cf, 4 9 Haemaphysalis 

leachi group) 
ex. Paradoxurus niger (Paradoxurus 


Hoshangabad District (2235'N, 7740'E), India 

C. A. Crump 
Received from Bombay Nat. Hist. Soc. 

K77 1914.11.17.16-35 

Haemaphysalis spinigera var. novae-guineae 
TYPE (19 cf, 4 9, 1 N Haemaphysalis novae- 
guineae paralectotypes) 
ex. Kangaroo 
Sattelberg, Huon Gulf (0710'S, 147'25'E), Papua 

New Guinea 

Prof. F. Forster 

Publications: Hirst (1914: 328), original descrip- 
tion; Nuttall & Warburton (1915: 452); Roberts 

K78 1914.11.17.1-15 

Haemaphysalis spinigera var. novae-guineae 

TYPE (1 cf Haemaphysalis novaeguineae lecto- 

type; 9 cf, 1 1 9 Haemaphysalis novaeguineae 

ex. Perameles sp. ? 
Sattelberg, Huon Gulf (07'10'S, 14725'E), Papua 

New Guinea 
Prof. F. Forster 

Publications: Hirst (1914: 328) original descrip- 
tion; Nuttall & Warburton (1915: 452); Roberts 
(1963: 57) selected lectotype and paralectotypes. 

K79 1914.11.17.36-46 

Haemaphysalis spinigera var. novae-guineae 
TYPE (6 cf, 10 9, 1 N Haemaphysalis novae- 
guineae paralectotypes) 
ex. Hydromys sp. 
Sattelberg, Huon Gulf (0710'S, 14725'E), Papua 

New Guinea 

Prof. F. Forster 

Publications: Hirst (1914: 328), original descrip- 
tion; Nuttall & Warburton (1915: 452); Roberts 
(1963: 57) designated paralectotypes. 

K80 1912.6.21.66 

1 cf Haemaphysalis parmata 

ex. Neotragus pygmaeus 

Bebaianiha (0543'N, 0027'W) or other 

populated places of this name nearby, Ghana 
H. G. F. Spurrell 

K81 None 

13 N Haemaphysalis spinigera 

ex. Lepussp. 

Hoshangabad District (2235'S, 7740'E), India 


C. A. Crump 

Received from Bombay Nat. Hist. Soc. 

Haemaphysalis spinigera is a vector of the viruses 

Kaisodi and Kyasanur forest disease. 

K82 1912.10.31.36-46 

Haemaphysalis sp. (28 cf, 7 9 Haemaphysalis 

ex. Wild cat 

Harar (09 1 9'N, 4209'E), Ethiopia 
6. 11.1912 
G. Kristensen 

K83 None 

Haemaphysalis leachi] (1 9 Haemaphysalis 

ex. Canis familiar is 

Himo River (0333'S, 3730'E), Tanzania 
Robin Kemp 

K84 19 12.6.21. 22-32 pt 

Haemaphysalis cinnabarina var. punctata (2 cf, 3 

9 Haemaphysalis sulcata) 
ex. Sheep 

Larnaca (3455'N, 3338'E), Cyprus 
C. B. Horsbrugh 

K85 19 12.6.21. 22-32 pt 

2 cf , 3 9 Haemaphysalis sulcata 

ex. Sheep 

Larnaca (3455'N, 3338'E), Cyprus 


C. B. Horsbrugh 

Note: A portion of this collection was originally 
entered under Haemaphysalis cinnabarina var. 

K86 None 

Haemaphysalis warburtonil (1 cf, 2 9, 2 N 

Haemaphysalis tibetensis TYPES 
1 cf (holotype) 2 9, 2 N (paratypes) 
ex. Canis familiar is 
Yatung (2725'N, 89 WE) (ca. 3896 m) mouth of 

Chumbi Valley, Tibet 

D. Macdonald 

Publication: Hoogstraal (1965: 452), original 

K87 1912.6.21.247-250 

Hyalomma syriacum (3 cf, 1 9 Hyalomma 


ex. Testudo graeca ibera 
Hammam Rirha (3626'N, 0228'E), Algeria 
24. V. 1911 
Hon. Walter Rothschild & Ernst Hartert 



K88 1912.6.21.251-253 

Hyalomma syriacum (Species epithet crossed out 

and aegyptium inserted) (4 9 Hyalomma 

anatolicum excavatum) 
Host unknown 
North West Morocco 
Received from Hon. W. Rothschild 

K89 1912.6.21.240-6 

Hyalomma aegyptium (2 cf, 1 9 Hyalomma 

Host unknown 

Hammam Rirha (3626'N, 0228'E), Algeria 

C. Hilgert 

Hyalomma marginatum is a vector of the viruses 
Dhori, West Nile, tickborne encephalitis, Bhanja, 
Crimean Congo hemorrhagic fever, Tamdy and 

K90 19 11. 12.9. 1-320 (pt) 

Hyalomma aegyptium (2 9 Hyalomma 


ex. Aged donkey (Equus asinus) 
Voi (0323'S, 38'34'E), Kenya 
Robin Kemp 

K91 None 

8 N Hyalomma sp. 

Host unknown 

Biskra (3450'N, 0540'E), Algeria 


Hon. L. W. Rothschild 

K92 19 19.8. 14. 1-70 (part) 

3 N Hyalomma sp. 

ex.? Dipodillus (gerbil) (Gerbillus (Dipodillus) sp.) 

('host similar to 108') 

Guelt-es-Stel (3509'N, 0302'E), Algeria 


Hon. Walter Rothschild and Karl Jordan 

K93 1912.6.21.202 

Rhipicephalus sp. (4 N Hyalomma sp.; 1 9 

Rhipicephalus pulchellus; 10 N Rhipicephalus 


ex. Lepus sp. 

Harar (09 1 9'N, 4209'E), Ethiopia 
G. Kristensen 

Rhipicephalus pulchellus is a vector of the 
viruses Dugbe, Crimean Congo hemorrhagic 
fever, and Nairobi sheep disease. 

K94 None 

Ixodes var. offecialis (2 9 Ixodes alluaudi) 

ex. Lophuromysjlavopunctatus zena 

Aberdare Mts (0025'S, 3638'E), Kenya 


Robin Kemp 

K95 None 

Ixodes var. offecialis ( 1 9 Ixodes alluaudi) 

ex. Lophuromys fJavopunctatus zena 

Aberdare Mts (0025'S, 3638'E), Kenya 1 1 ,000 ft 

(3,353 m) 
Robin Kemp 

K96 None 

Ixodes tenuirostris (2 9 Ixodes alluaudi) 

ex. Otomys tropicalis elgonis 

Mt Ilkinangop (0038'S, 3642'E), Aberdare Mts, 

Robin Kemp 

K97 None 

Ixodes tenuirostris (1 9 Ixodes alluaudi) 

Host unknown 

Rombo, Kilimanjaro (0305'S, 3720'E), 

Robin Kemp 
Note: This 9 has been previously pinned. 

K98 None 

[undetermined genus & species] (1 N Ixodes 

ex. Mus jacksoni (Possibly Anomalurus 

derb ianus Jackson i) 
Mutaragwa, Aberdare Mts (locality not verified*), 

R. Kemp 

*There is a Mutarakua Hill (0047'S, 3514'E) in 
the Aberdares that is a possible locality for this 

K99 1912.6.21.280-2 

4 9 Ixodes cordifer 

ex. Pseudoircornia (probably Pseudocheirus sp.) 

Sattelberg, Huon Gulf (0710'S, 14725'E), Papua 

New Guinea 
Prof. F. Forster 

K100 1912.6.21.283 

1 9 Ixodes cordifer 

ex. Dactylopsida trivirgata 

Sattelberg, Huon Gulf (0710'S, 14725'E), Papua 

New Guinea 
Prof. F. Forster 

Note: label in vial seen by Prof. R. A. Cooley 
1946. 3 slides of dissections (RML 22285). 

K101 1912.6.21.284-8 

4 9 Ixodes cordifer 

ex. Phalanger maculatus 

Sattelberg, Huon Gulf (0710'S, 147'25'E), Papua 

New Guinea 
Prof. F. Forster 



K102 1912.6.21.279 

1 d 1 Ixodes cordifer 

ex. Perameles sp. 

Sattelberg, Huon Gulf (0710'S, 14725'E), Papua 

New Guinea 
Prof. F. Forster 

K103 1912.12.4.11 

1 9, 4 N Ixodes fecialis 

cx Fclis so 

Studley Park, Melbourne (3749'S, 14458'E), 

A. Coles 

Note: Reverse of the vial label bears the 
inscription Native cat. 

K104 1912.6.21.254 6 

5 9, 2 N Ixodes hexagonus 

ex. Mustela vulgaris (Mustela nivalis) 

Tring (5 1 47'N, 0039'W), England 


J. F. Cox 

K105 1912.6.21.259-61 

1 9, 2 N, 7 L Ixodes hexagonus (1 9, 2 N Ixodes 

ex. Mustela erminea 
Tring (5 1 47'N, 0039'W), England 
J. F. Cox 

K106 1912.6.21.262-3 

4 N Ixodes hexagonus (4 N, 8 L Ixodes 


ex. Meles taxus (Meles meles) 
Senj (45WN, 1454'E), Yugoslavia 
F. Dobiasch 

K107 None 

22 N, 6 L Ixodes hexagonus 

ex. Mustela erminea 

Loch Tay (5628'N, 04 1 8'W), Lawes, Scotland 


L. G. Esson 


1 9 Ixodes hexagonus 
Host unknown 
Faugeres*, S. France 

W. Davy 

N. C. Rothschild 

*Three possible localities for Faugeres, France: 

4334'N, 03'11'E; 4428'N, 0358'E; 4458'N, 


K 109 None 

2 N Ixodes hexagonus 
ex. Mustela putorius 

Carmarthen (5 152'N, 04 1 8'W), Wales 


F. W. Frohawk 

Kl 10 1919.8. 14.1-70 (part) 

1 9 Ixodes hexagonus 

ex. dog Canis familiaris 

Bridgnorth (5232'N, 0225'W), England 


Miss Frances Pitt 

Kill 1912.12.4.9-10 

1 rf, 1 9 Ixodes loricatus 

ex. Didelphis marsupialis aurita 

Mar de Espanha (2052'S, 43'00'W), Estado de 

Minas Gerais, Brazil 
Z. F. Zikan 

K112 1919.8. 14.1-70 pt 

2 <?, 5 9 Ixodes loricatus 

ex. Lutreolina crassicaudata 

Parana Delta (vicinity of 3343'S, 5915'W), 

Robin Kemp 

K113 1912.6.21.289-292 

Ixodes loricatus var. spinosus (4 d 1 Ixodes luciae) 

ex. Didelphis marsupialis 

San Esteban (1 1'28'N, 69'22'W), Venezuela 


Kl 14 None 

4 9, 1 5 N, 1 L Ixodes ornithorhynchi 

ex. Ornithorhynchus paradoxus (Ornithor- 

hynchus anatinus) 
Tyalgum Tereed River, perhaps Tyalgum Creek 

(2822'S, 153'13'E), New South Wales, 

Received from W. F. H. Rosenberg 

K115 1912.6.21.272-3 

Ixodes rasus (2 9 Ixodes cumulatimpunctatus) 

ex. Cricetomys gambianus 

Bibianiha (0543'N, 0027'W) or other populated 

places of this name nearby, Ghana 
H. G. F. Spurrell 

K116 1912.6.21.264 

3 9 Ixodes ricinus (2 9 Ixodes ricinus) 
ex. Canis familiaris 

Porlock Weir (5 1'12'N, 0340'W), England 


F. J. Cox 

Ixodes ricinus is a vector of the viruses Tettnang, 

Tribec, Louping ill, tickborne encephalitis, West 

Nile, Crimean Congo hemorrhagic fever, 

Lipovnic, Uukuniemi, Sumakh, Eyack and 




Kl 17 None 

Ixodes tasmani! (2 cf, 1 9, 1 N Ixodes tasmani) 

ex. Nest of Pseudochirus cooki (Pseudocheirus 


N. W. Coast, Tasmania, Australia 
R. N. Atkinson 

K118 1912.10.31.7-10 

Ixodes tenuirostris (1 cf, 2 9, 3 N Ixodes 


ex. Microtus glareolus (Clethrionomys glareolus) 
Braunton (5 1 06'N, 04 1 1 'W), England 
W. Holland 

K119 1912.6.21.267 

1 N Ixodes vespertilionis 

ex. Rhinolophusferrumequinum 

Cave de Meaunes, Southern France (locality not 

Dr K. Jordan 

K120 1912.6.21.268-9 

4 N, 1 L Ixodes vespertilionis 

ex. Bat? 

Monchique (3720'N, 0829'W), Portugal 


Dr K. Jordan 

K121 1912.10.31.11 

1 9 Ixodes vespertilionis 

ex. Rhinolophusferrumequinum 

Braunton (5 1 06'N, 04 1 1 'W), England 


W. Holland 

N. C. Rothschild 

Note: This 9 is in alcohol but there is a pin 

through the idiosoma. 

K122 1912.6.21.266 

2 N Ixodes vespertilionis 

ex. Rhinolophusferrumequinum 

St Genes de Malgoires (Saint-Genies-de- 

Malgoires) (4357'N, 04 13'E), France 
Albert Hugues 


1 N Ixodes vespertilionis 

ex. bat 

Oumasch, Algeria (locality not verified) 


Hon. Walter Rothschild & Ernst Hartert 

Note: Theodor (1967: 94) cited a collection of 2 cf , 

3 9 Nycteribia schmidlii schmidlii off Plecotus 
auritus christiei, Oumasch near Biskra, Algeria, 
5. III. 1911, Hon. W. Rothschild and E. Hartert. 
Quite probably this bat was host for both species 
of ectoparasite. 

K124 1912.6.21.270 

1 9 Ixodes vespertilionis 

ex. Rhinolophus hipposideros 

Uj-Moldova (4444'N, 2 1 42'E), Romania 


Lintia Dionisius 

K 125 None 

9 N, 5 L Rhipicephalus h. haemaphysaloides (4 N, 

2 L Rhipicephalus haemaphysaloides) 
ex. Lepussp. 

Hoshangabad(2246'N, 7745'W), India 

C. A. Crump 
Received from Bombay Nat. Hist. Soc. 

K126 1912.6.21.203-204 

Rhipicephalus kochi(\ d 1 Rhipicephalus jeanneli) 

ex. grass in tent 

Voi (0323'S, 3824'E), Kenya 

R. Kemp 



Rhipicephalus sp. (3 c? Rhipicephalus kochi; 1 cf 
Rhipicephalus longicoxatus; 6 cf, 1 9 Rhipi- 
cephalus pravus) 

ex. Aged donkey (Equus asinus) 

Voi (0323'S, 3834'E), Kenya 

29. IV.1910 

Robin Kemp 

Rhipicephalus pravus is a vector of Kadam 


K128 1912.6.21.239 

Rhipicephalus sp. (1 9 Rhipicephalus kochi group) 
ex. Ox 

Curepipe(2019'S, 57'31'E), Mauritius 
3. XI. 1911 
C. Brishoe 

Note: Tentative identification: Engorged mis- 
shapen specimen. 

K 129 None 

1 cf, 1 9, 10 N Rhipicephalus pravus 

Host unknown 

Taveta (0325'S, 3743'E), Kenya 


Robin Kemp 

K130 1953.10.9.17 

1 rf Rhipicephalus pravus (1 cf, 1 N Rhipicephalus 


ex. Jerboa? (Jaculus sp. ?) 
Voi (0323'S, 3834'E), Kenya 
22. IV. 1910 
Robin Kemp 

K131 1911. 12.9. 1-320 (part) 

Rhipicephalus sp. (25 rf, 27 9 Rhipicephalus 

Host unknown 



9 Rhipicephalus 

Rombo, Kilimanjaro (0305'S, 3720'E), 

15. VI. 1910 
Robin Kemp 

K132 1912.6.21.229 

1 cf, 1 9 Rhipicephalus pulchellus 

ex. Homo sapiens off ground in tent 

Nairobi (0 1 1 7'S, 3649'E), Kenya 


Robin Kemp 

K133 1919.8. 14.1-70 pt 
Rhipicephalus simus (1 c 


ex. Vulpes vulgaris ( Vulpes vulpes) 
Toscana(4325'N, 1 POO'E), Italy 

N. Cimballi 

Rhipicephalus sanguineus is a vector of Wad 
Medani virus and the rickettsia of boutonneuse 

K134 1919.8. 14.1-70 pt 

Rhipicephalus simus (1 9 Rhipicephalus 

Host unknown 

Firenze (4346'N, 1 1 1 5'E), Italy 
N. Cimballi 

K135 1912.12.4.18 

Rhipicentor ( 1 9 Rhipicephalus sanguineus) 

ex. dog (Canis familiar is) 

Livingstone ( 1 220'S, 30 1 8'E), Zambia 


A. Douglas 

K136 1912.6.21.231-232 

2 9 Rhipicephalus sanguineus 

ex. Canis familiaris 

Dehra Dun (3019'N, 78'04'W), India 


Mrs R. Parson 

K137 1912.6.21.234-38 

6cf, 149 1 N Rhipicephalus sanguineus 

ex. Canis familiaris 

Manaus (03 1 0'S, 60WW), Brazil 


Rev. A. Miles Moss 


Rhipicephalus sp. (1 9 Rhipicephalus sanguineus) 

ex. Gerbillus indicus (Gerbillus sp.) 

Khandala (1 855'N, 7325'E), India 


J. Assmuth 

Note: Tentative identification: Teneral runt 


K139 1912.6.21.205-220 
Rhipicephalus sp. (17 

6 9 Rhipicephalus 

ex. Lepus sp. 

Mountains near Droh (3452'N, 0553'E), Algeria 


Hon. Walter Rothschild & Ernst Hartert 

K140 1912.6.21.201 

Rhipicephalus sp. (1 9 Rhipicephalus sanguineus) 

ex. Erinaceus europaeus 

Saint-Genies-de-Malgoires (4357'N, 0413'E), 

21. III. 1909 
A. Hugues 

K141 19 19.8. 14. 1-70 part 

Rhipicephalus sp. (2 9 Rhipicephalus sanguineus) 

ex. hedgehog 

Hammam Meskoutine (3627'N, 0716'E), 

Hon. Walter Rothschild & Karl Jordan 

K142 1912.6.21.230 

Rhipicephalus sp. (1 d 1 Rhipicephalus simus) 

ex. Vulture 

Mduna River, Hlabisa District (2810'S, 32'10'E), 

Zululand Region, Republic of South Africa 
F. Toppin 

Received from Natal Museum, Pietermaritzberg 
Rhipicephalus simus is a vector of the viruses 
Thogoto and Nairobi sheep disease. 

K143 None 

1 cf, 5 9 Rhipicephalus simus 

Host unknown 


Dr E. J. Baxter 

K 144 None 

1 N Rhipicephalus sp. 
Host unknown 

Biskra (3450'N, 0540'E), Algeria 


2 N Rhipicephalus sp. 

ex. Elephantulus rufescens 
Voi (03'23'S, 3834'E), Kenya 
Robin Kemp 

K 146 None 

10 N, 2 L Rhipicephalus sp. 

ex. Elephantulus rufescens 

Voi (0323'S, 3834'E), Kenya 


Robin Kemp 


2 N Rhipicephalus sp. 

ex. Elephantulus rufescens 

Voi (0323'S, 3834'E), Kenya 


Robin Kemp 


Rothschild specimens missing from British Museum (Natural History) collection 

K148 1912.6.21.3-20 
Argas persicus 
ex. fowls 
Larnaca, Cyprus 

K149 1911. 12.9.1-320 pt 
Amblyomma gemma 
Host unknown 
Kilimanjaro, Brit. E. Africa 

K150 1912.6.21.41 1-415 
Aponomma gervaisi var. trimaculatus 
ex. Iguana 
Kelsey Creek, N. Queensland 

K151 1919.8. 14.1-70(pt) 

Haemaphysalis leachi 

ex. Hedgehog 

Meskoutine Hamman, Mespot. 

K152 1912.10.31.36-46 
Haemaphysalis leachi indica 

ex. Wildcat 
Harar, Abyssinia 

K153 1911. 12.9.1-320 pt. 
Hyalomma truncatum 
ex. Donkey 
British East Africa 

K154 1912.6.21.265 
1 9 Ixodes ricinus 
Host unknown 
Hamman Rirha, Algeria 

K155 1919.8. 14.1-70 pt. 
Ixodes vespertilionis 
ex. Vespertilio euryalis 

K156 1912.6.21.188-200 
Rhipicephalus sanguineus 
ex. Erinaceus europaeus 
Montchique [ = Monchique], Portugal 

Locality list for ticks in the Rothschild collection 

N = Nuttall, K = British Museum (Natural History) collections. Numbers in parentheses indicate 
missing collections. 

Abyssinia (Kl 52) 

Algeria N561, N3537, N3538, N3539, (N3536), Kl, K2, K87, K89, K91, K92, K123, K139, K141, 

Argentina N588, K19, K20, K21, K22, K34, Kl 12 
Australia N546, N547, N550, N552, N553, N556, N567, N569, N633, N642, N643, N645, N646, 

N650, N669, Nl 162, N121 1, N1212, N1213, N1392, N1393, N1397, (N551), (N571), (N644), K6, 

K33, K40, K41, K42, K43, K44, K45, K46, K47, K49, K57, K58, K59, K60, K103, Kl 14, Kl 17, 


Belgium N597,(N631) 

Bolivia N560, N562, N566 

Brazil N641,(N544), K8, K10, K17, K18, K26, K27, K28, K29, K30, K31, K32, Kl 1 1, K137 

British East Africa (K 1 49), (K 1 53) 

Canada N1105, N1387, N1390, N1399, N1497, N1498, N1499, N1500, N1501, N1502, N1503, 

N 1 504, N3502, (N596), (N 1 388) 
Chile N 1389 
Christmas Island, N360 
Colombia K5, K24 
Cyprus K84, K85,(K148) 

Ecuador K23 

Egypt (N539) 

England N580, N593, N598, N599, N600, N601, N602, N603, N604, N605, N607, N610, N611, 
N618, N621, N622, N623, N652, N1067, N1068, Nl 152, Nl 159, Nl 160, Nl 163, N3160, (N216), 
(N217), (N298), (N322), (N324), (N395), (N396), (N509), (N518), (N656), K104, K105, K110, 

Ethiopia K36, K37, K38, K63, K64, K65, K66, K82, K93 


France N523, K55, K108, Kl 19, K122, K140 

Galapagos Islands N397 
Germany, Democratic Republic of K3 
Germany, Federal Republic of N608 
Ghana Kl 1, K48, K80, Kl 15 
Guyana K15,K16 

India N572, N573, N576, N581, N582, N583, N592, N892, (N3158), (N3159), K76, K81, K125, 

K136, K138 

Indonesia N72, N73, N74 
Ireland N61 2, (N630) 
Italy K133,K134 

Jamaica K9 
Japan N579 

Kalao Island (N76) 
Kalas Island (N93) 

Kenya K12, K13, K61, K75, K90, K94, K95, K96, K98, K126, K127, K128, K129, K130, K132, 

Liberia K39, K67 

Louisade Archipelago Nl 166 

Madagascar, Democratic Republic of N670 

Malawi N563 

Mauritius K127 

Mexico N557, N568, N640, N647, N648, N1386, N1394, N1396, K25 

Morocco K88 

Papua New Guinea K51, K77, K78, K79, K99, K100, K101, K102 
Paraguay N 1391 
Philippine Islands N545 
Portugal Kl 20, (K 156) 

Romania Kl 25 

Sardinia Nil 51 

Scotland N667, K 107 

Senegal N888 (W. Rothschild) 

South Africa, Republic of N649, Nl 167, Nl 167a, K50, K142 


Switzerland N619, N624, N625, N1069, Nl 161 

Tanzania K52, K62, K68, K69, K70, K71, K72, K73, K74, K83, K97, K131, K143 
Tibet K86 
Trinidad-Tobago K53 

Uganda-Zaire border N559 

Unknown N549, N554, N555, N558a, N558b, N564, N565, N570, N574, N575, N575x, N584, N585, 
N589, N590, N591, N613, N617, N634, N637, N638, N639, N826, N1066, Nl 153, Nl 154, Nl 164, 
N1395, N1398, N3138, N3330, (N75), (N77), (N78), (N90), (N91), (N92), (N516), (N517), (N548), 
(N586), (N587), (N595), (N620), (N628), (N629), (N632), (N674), (N3136), (N3139), (N3140), 
(N3141),(N3157),(N3351), K14 

U.S.A. N594, N626, N666, Nl 399, K4, K7 

U.S.S.R. K56 

Venezuela Kl 13 

Wales N606, N609, (N746), K109 

Yugoslavia K 106 

Zambia K 135 

Zimbabwe N577,N578 



Host list for ticks in the Rothschild collection 

Numbers in parentheses indicate missing collections. All information relating to missing collections is 
taken directly from Nuttall's catalogue or British Museum (Nat. Hist.) collection cards. 


Anaconda (N3 141) 

Anomalurus derbianusjacksoni K98 

Anteater K24 

Armadillo K34 

Arvicola terrestris N662 

Ateles geoffroyi geoffroyi N565, N640 

BatN591,K120, K123 

Bettongia lesueri N646 

Bettongia penicillata N 1 393 

Bird (N397) 

Blastoceros bezoarcticus K2 1 

Boa (N548) 

Bradypus tridactylus (N544), (N335 1 ) 

Camel (N586) 

Canis familiaris N572, N573, N576, N579, 

N645, N892, K9, K38, K56, K60, K83, K86, 

K110, K116, K135,K136, K137 
Canis, sp. K19 
Canis vulpes (N2 1 7) 
Cat, black N504 
Cat, wild K82 
Cattle K9 

Cercartetus concinnus N 1 1 62 
Cervus capreolus (N509) 
Chicken K4 

Clethrionomys glareolus N6 1 9, K 1 1 8 
Coendou bicolor simonsi N560 
Coendou prehensilis K 1 8 
Columba livia K3 
Cricetomys gambianus Kl 1 5 

Dactylopsida trivirgata K 1 00 

Damaliscus lunatus N 1 1 67, N 1 1 67a 

Dasyurus geoffroyi N650 

Deer K53 

Didelphis marsupialis N64 1 , Kl 1 3 

Didelphis marsupialis aurita Kill 

Didelphis sp. N638, N647, N648, N1391 

Diemenia superciliosa (N57 1 ) 

Diemenia textilis N569, N642 

Diomedea salvinii (N5 1 6) 

Dog, 'Badlington' K67 

Donkey (Kl 53) 

Duck, wild N6 17 

Dusicyon gymnocerus K22 

Echidnas?. N3138 

EirabarbaraKll, K27 

Elephantulus rufescens K145, K146, K147 

Equusasinus K14, K90, K127 


Erinaceus albiventris (N539) 

Erinaceus europaeus N597, N599, N608, (N324), 

(N631), K139,(K156) 
Erithacm rubecula (N396) 

Feliscatus K65 
Pel is concolor N637 
Felis serval K68 

Fowls (Kl 48) 
Fox (N632) 
Fox, chubut K20 
Fratercula arctica N634 

Genetta stuhlmanni K75 

Gerbillus campestris N3537, N3538, N3539 

Gerbillus (Dipodillus) sp. K92 

Gerbillus gerbillus N 1 398 

Giraffa camelopardalis N558a, N558b, N888 (W. 

Grison vittata K28 

Hedgehog K141,(K1 51) 

Homo sapiens N1498, K9, K39, K132 

Hyaena hyaena N575X, N1395, K64 

Hydrochoerus hydrochaeris K23 

Hydromys sp. K79 

Hystrix sp. K50 

Ictonyx striatus N577 
Iguana (Kl 50) 
Isoodon macrourus N669 
Isoodon obesulus N 1 2 1 3 

Jaculussp. K130 

Kangaroo N546, N552, N553,(N551), K77 

Lagorchestes conspicillatus N 1 392 
Lagostrophusfaciatus K33 
Lepussp. K81, K93, K125, K139 
Lophuromys flavopunctatus zena K94, K95 

Lutra vulgaris (N2 1 6) 
Lutreolina crassicaudata Kl 12 

Macropus dorsalis K58 

Macropus fuliginosus N 1 397 

Macropus sp. N633 (N644) 

Manissp. Kl 1 

Manis tricuspsis N564 

Marten K66 

Megaderma cor N 1 1 64 

Meles meles K 1 06 

Microtus agrestis N623 

Microtus arvalis N624, N625, N 1069 

Monitor (N76),(N93) 



Monitor, Australian N550 

Monitor strix N554 

Morelia spilotes variegata N567 

Mus escularis N 1 1 66 

Mus minutus (N620) 

Mustela erminea N602, N603, N607, N610, 

N611, N612, N613, Ml 159, (N656), K105, 


Mustela nivalis N604, K104 
Mustela putorius N600, N606, N609 (N322), 


Mustela vison N1399 
Myocastor coypus N826 
Myopotomus coypu (N674) 
Myotis tricolor N649 
Myrmecophaga tridactyla K25 

Nasua nasua K26, K29, K3 1 , K32 
Neotoma alleni vetula N 1 394 
Neotragus pygmaeus K80 
Nestlings N594 
Notechis scutatus K49 

Ochotona princeps Nl 387 

Odocoileus virginianus N626, N666, Nl 386 

Ophiophagus bungaris (N90) 

Ornithorhynchus anatinus Kl 14 

Oryctolagus sp. K72 

Oryzomys longicaudatus N 1 389 

Otomys tropicalis elgonis K96 


OxenK12, K13 

Paradoxurus hermaphroditus K76 

Passer montanus N523 

Perameles sp. K78, K102 

Phalacrocorax aristotelis N652, N667 

Phalacrocorax graculus (N395) 

Phalanger maculatus K101 

Pipistrellus pipistrellus N593, Nil 51, Nil 52, 


'Piquoti' N639 

Plecotus auritus christiei Kl 23 
Polecat (N746) 
Procyon cancrivorus K30 
Pseudocheirus sp. Kl 1 7 
Pseudocheirus sp. probably K98 
Puffin (N5 18) 

Putorius energermanis (N 1 388) 
Putorius erminea (N298) 
Python sp. K5 1 

Rangifer tarandus N1500, N1501, N1502, 


Rattus macleari N360 
Raven K36 
Rhinoceros N563 

Rhinolophus ferrumequinum Kl 19, K121, K122 
Rhinolophus hipposideros K124 
Riparia riparia N605, N1068 

Sciurus niger K7 

Sciurus sp. N 1 396 

Sciurus vulgaris (N630) 

Scotophilus temmincki wroughtoni N592 

Sheep K35, K54, K84, K85 

Sminthopsis murina N 1 2 1 1 

Snake, black and white N78 

Snake, large N569 

Sorex araneus N62 1 

Speothos venaticus N562, N566 

Squirrel, flying N581 


Suricata suricata N578 

Sus verrucosus celebensis N72 

Sus verrucosus subsp. N73, N74 

Tachyglossus aculeatus N547, K40, K41, K42, 


Talpa sp. N583 
Tapirussp. K16 
Tapir us terrestris K10 
Tayassusp. N557 
Tenerec ecaudatus N670 
Testudo graeca ibera K87 
Testudo pardalis (N77), (N3 140) 
Tortoise N549, (N75), (N3 1 39) 
Tortoise, Algerian N561 
Tortoise, leopard N570 
Trachysaurus rugosus K6 
Turtle Kl 5 

Uria lachrymans (N5 1 7) 

Varanus civitatus (N9 1 ), (N92) 
Varanus sp. K44, K45, K46, K47, K48 
Vesper tilio euryalis (Kl 55) 
Vulpes vulpes N1067, Nl 160, K133 
Vulture K142 

Wallabia bicolor K58 
Wallabia sp. K59 

Zebra (N3 157) 
Zorilla N577 


I am most grateful to the Misses B. E. Brewster, formerly Arachnida Section, and P. D. Jenkins, 
Mammal Section, Zoology Department, British Museum (Natural History), and to Mr D. Macfarlane, 
Commonwealth Institute of Entomology, for assisting me in obtaining correct host designations and 
locality coordinates. I also thank Dr K. C. Kim, The Pennsylvania State University, for determination 
of the Anoplura. 



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Arthur, D. R. 1953. The systematic status of Ixodes percavatus var. rothschildi Nuttall & Warburton, 

1911. Parasitology, Cambridge 43 : 222-226. 

1963. British Ticks. Butterworth & Co. Ltd., London. i-ix, + 213 p. 
Cooley, R. A. & Kohls, G. M. 1945. The genus Ixodes in North America. Natn. Inst. Hlth. Bull. 184. 

246 p. 
Hirst, S. 1914. Report on the Arachnida and Myriopoda collected by the British Ornithologists' Union 

Expedition and the Wollaston Expedition in Dutch New Guinea. Trans, zool. Soc. Lond. 

20 : 325334. 
1916. On a new variety of European tick (Dermacentor reticulatus, var. aulicus, var. nov.). Ann. 

Mag. nat. Hist. (8), 17 : 308. 
Hoogstraal, H. 1953. Ticks (Ixodoidea) of the Malagasy faunal region (excepting the Seychelles). 

Their origins and host-relationships; with descriptions of five new Haemaphysalis species. Bull. Mus. 

comp. Zool. Harv. Ill : 37-1 13. 
1955. Notes on African Haemaphysalis ticks. I. The Mediterranean-littoral hedgehog parasite H. 

erinacei Pavesi, 1884 (Ixodoidea, Ixodidae). J. Parasit. 41 : 221-233. 
1956. African Ixodoidea. I. Ticks of the Sudan. Dep. Navy, Bur. Med. Surg.; Washington, D.C. 

1101 p. 
1965. Haemaphysalis tibetensis sp. n., and its significance in elucidating phylogenetic patterns in 

the genus (Ixodoidea, Ixodidae). J. Parasit. 51 : 452-459. 
, Carney, W. P., Kadarson, S. & Van Peenen, P. F. D. 1973. Haemaphysalis (Kaiseriana) 

celebensis Hoogstraal, Trapido, and Kohls (Ixodoidea, Ixodidae), a Wallacean member of the 

hystricis group: Identity, distribution, hosts, and ecology. /. Parasit. 59 : 556-562. 
, Trapido, H. & Kohls, G. M. 1965a. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, 

Ixodidae). The identity, distribution, and hosts of H. (Kaiseriana) hystricis Supino. J. Parasit. 

51 : 467^80. 

& 19656. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, Ixodidae). H. 

(Kaiseriana) celebensis sp. n., from a wild boar in Celebes. J. Parasit. 51 : 1001-1003. 
Hopkins, G. H. E. & Rothschild, M. 1953-1971 . An Illustrated Catalogue of the Rothschild Collection 

of Fleas (Siphonaptera) in the British Museum (Natural History). Vols. I-V. British Museum 

(Natural History), London. 
Kaufman, T. S. 1972. A revision of the genus Aponomma Neumann, 1899 (Acarina: Ixodidae). Ph.D. 

thesis University of Maryland, 289 pp. 

Neumann, L. G. 1904. Notes sur les Ixodides. II. Archs. Parasit. 8 : 444^64. 
Nuttall, G. H. F. 1910. New species of ticks (Ixodes, Amblyomma, Rhipicephalus). Parasitology, 

Cambridge 3 : 408^1 16. 
& Warburton C. 1911. Ixodidae. Section II. The genus Ixodes. In Nuttall el ai, Ticks. A 

monograph of the Ixodoidea. Part II: 1 33-293. Cambridge University Press, Cambridge. 

& 1915. Ixodidae. The genus Haemaphysalis. In Nuttall el /., Ticks. A monograph of the 

Ixodoidea. Part III: 349-550. Cambridge University Press, Cambridge. 
Roberts, F. H. S. 1960. A systematic study of the Australian species of the genus Ixodes (Acarina: 

Ixodidae). Aust. J. Zool. 8 : 392-485. 
1963. A systematic study of the Australian species of the genus Haemaphysalis Koch (Acarina: 

Ixodidae). Aust. J. Zool. 11 : 35-80. 

1970. Australian Ticks. Commonwealth Scientific and Industrial Research Organization, 

Australia; Melbourne, i-iv, 267 p. 
Robinson, L. E. 1926. The genus Amblyomma. In Nuttall el ai, Ticks. A monograph of the Ixodoidea. 

Part IV: 1 302. Cambridge University Press, Cambridge. 
Taylor, F. H. 1946. Spiders, ticks and mites including the species harmful to man in Australia and New 

Guinea. Section 1 . Descriptive. Serv. Publs. Sch. publ. Hlth. trop. Med. Univ., Sydney. (6): 7-234. 
Theodor, O. 1967. An illustrated Catalogue of the Rothschild Collection of Nycteribiidae (Diptera) in 

the British Museum (Natural History). London-British Museum (Natural History), Publication No. 

655. pp. viii + 506 + 5 plates. 
Warburton, C. & Nuttall, G. H. F. 1909. On new species of Ixodoidea, with a note on abnormalities 

observed in ticks. Parasitology, Cambridge! : 57-76. 
/ u nipt. F. 1952. The ticks of sea birds. Aust. Nat. Antarctic Res. Rep. Series B. 1 : 12-20. 

British Museum (Natural History) 

An Illustrated Catalogue of the 
Rothschild Collection of Fleas 
(Siphonapterd) in the British Museum 
(Natural History) 

With keys and short descriptions for the 
identification of families, genera, species 
and subspecies of the Order 

The collection of fleas in the British Museum (Natural History) ranks among the most important 
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Rothschild and presented by him to the Nation in 1913 with the proviso that a catalogue be 
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which some 90% of the known 2,000 or so species are represented. It is a work of 
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This series of volumes provides a comprehensive taxonomic monograph on the group, with 
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excellent drawings. 

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by G. H. E. Hopkins and M. Rothschild 

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general collections of the British Museum (Natural History). 

By James E. Keirans 

Hydroids and medusae of the family Campanulariidae recorded 
from the eastern North Atlantic, with a world synopsis of genera. 
By P. F. S. Cornelius 


Printed by Henry Ling Ltd, Dorchester 

Bulletin of the 

British Museum (Natural History) 

Hydroids and medusae of the family 
Campanulariidae recorded from the 
eastern North Atlantic, with a world 
synopsis of genera 

P. F. S. Cornelius 

Zoology series Vol 42 No 2 29 April 1982 

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Vol 42 No 2 pp 37-148 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 29 April 1982 

I * 

Hydroids and medusae of the family \* UBRA V ^ 

Campanulariidae recorded from the eastern North ^/ 
Atlantic, with a world synopsis of genera 

P. F. S. Cornelius 

Zoology Department, British Museum (Natural History), Cromwell Road, London 
SW7 5BD, England 


Synopsis 38 

Type designations 38 

Introduction 38 

Taxonomy and the phenotype 39 

Ecology 43 

Geographical distribution 44 

Key to species 45 

Taxonomic section 47 

Family Campanulariidae 47 

The subfamily divisions and their nomenclature 49 

Subfamily Campanulariinae 50 

Genus Campanularia 51 

Campanularia crenata 52 

Campanularia hincksii 53 

Campanularia volubilis . . 55 

Genus Orthopyxis 57 

Orthopyxis crenata 58 

Orthopyxis integra 60 

Genus Rhizocaulus 67 

Rhizocaulus verticillatus 67 

Subfamily Clytiinae nom. nov 69 

Genus Clytia 70 

Clytia discoida 

Clytia hemisphaerica 73 

Clytia hummelincki 82 

Clytia islandica 84 

Clytia linearis 84 

Clytia mccradyi 87 

Clytia paulensis . 88 

Clytia incertae sedis . 91 

Subfamily Obeliinae 

Genus Gonothyraea 91 

Gonothyraea loveni 92 

Genus Hartlaubella 94 

Hartlaubella gelatinosa 

Genus Laomedea 

Laomedea angulata. 98 

Laomedea calceolifera 102 

Laomedea flexuosa 105 

Laomedea neglecta 107 

Laomedea pseudodichotoma Ill 

Bull. Br. Mus. nat. Hist. (Zool.) 42 (2) ; 37-148 

Issued 29 April 1982 



Genus Obelia 

Obelia bidentata . . . . . 

Obelia dichotoma . . . . . 

Obelia geniculata . 

Obelia spp. indet 120 

Taxa now referred to other families 




References 126 

Index 14 


Hydroids and medusae of the family Campanulariidae recorded from the eastern North Atlantic are 
revised. Of more than three hundred nominal species just twenty-three are regarded valid. Their 
taxonomy, nomenclature, morphology, ecology, reproduction, distribution and relationships are 
discussed and a key is provided for their identification. The genera occurring in the eastern North 
Atlantic are revised in detail. In addition the genera not represented in this area are briefly reviewed and 
notes are included to update a previous revision of the genus Obelia. 

Type designations 

Type material is designated of the nominal species Campanularia laevis Couch, 1844, to 
facilitate comparison with Campanularia laevis Hartlaub, 1905, for which a new name is 
proposed (p. 54). Lectotype material is designated ofSertularia uniflora Pallas, 1766 (p. 78), 
Laomedea gracilis Sars, 1850 (for which a new name is proposed, p. 78), and Campanularia 
intertexta Couch, 1844 (p. 122). 

Type species are designated of two genera and a subgenus: Cymodocea Lamouroux, 1816, 
family Plumulariidae (p. 121); Eucope Gegenbaur, 1856 (p. 71); subgenus Eucampanularia 
Broch, 1910 (p. 52). The type genus of the subfamily Clytiinae nom. nov. is designated as 
Clytia Lamouroux, 1812 (p. 69), and that of the subfamily Phialiinae (Family Lovenellidae) 
asPhialium Haeckel, 1879 (p. 69). 


The family Campanulariidae is represented in all oceans. The species are found mainly 
within continental shelf depths, and some occur intertidally. Many of the genera and species 
are cosmopolitan. For example, many of the species recorded from New Zealand by Ralph 
(1957) occur also in British waters (see Geographical distribution, p. 44). 

Several European authors have revised the species of Campanulariidae recorded from 
their home waters (e.g. Hincks, 1868;Linko, 1911; Broch, 1918;Nobre, 1931;Kramp, 1935; 
da Cunha, 1944; Vervoort, 1946a; Leloup, 1947; Russell, 1953; Naumov, 1960, 1969; 
Patriti, 1970; Rossi, 1971). But apart from the synoptic lists of Bedot (1901, 1905, 1910, 
1912, 1916, 1918, 1925) and a review of the genus Obelia (by Cornelius, 1975a) there has 
been no previous attempt to compare the nominal species recorded from the whole of the 
eastern North Atlantic. As a result many straightforward taxonomic questions have been left 
unresolved. No doubt the wide range of phenotypic variation typical of this family has 
caused problems. But it is apparent also that the wide geographical ranges of many of the 
species has led to unwitting duplication of work between countries. The resulting literature is 
widely scattered, and this too has imposed problems since only a few libraries have all the 
relevant publications. 

This report attempts to revise the species of the family Campanulariidae, excepting those 
in the genus Obelia, recorded from the eastern North Atlantic. Obelia is excluded since it 
was revised recently (Cornelius, 1975a) but notes are included to update that revision. The 



area extends southwards to the equator and west to the 30 meridian. It includes the 
Mediterranean, Black and Baltic Seas, and extends in one region beyond 30 W to include the 
coastal waters of Greenland. In general there have been few records of the family north of 
80 N, but this must reflect collecting difficulties since several species have been reported 
from these high latitudes (Linko, 1911). One dubious tropical species, Orthopyxis africana, is 
not treated fully because it has been recorded only once, near the southern limit of the area. 

Most genera of Campanulariidae are nearly cosmopolitan. It happens that the majority are 
represented in the eastern North Atlantic. Further, most were described first from European 
waters; so that a revision of the NE Atlantic genera comes close to a complete revision. A 
comprehensive generic synopsis is not intended here, however, since not all the nominal 
species are covered. But at least an interim generic list can be given (Table 1). Most of the 
nominal genera are discussed, whether or not they have been recorded in the NE Atlantic, 
and the relevant discussion sections can be found through the index. All are at least 

Table 1 The subfamilies and accepted genera of the Campanulariidae. Those not recorded from the 
NE Atlantic are treated only briefly in this paper, and are marked by an asterisk. 


Recent redefinition, if any 



Campanularia Lamarck, 1816 
*Eucalix Stechow 192 la 
*Orthonia Stechow, 1923a 
Orthopyxis Agassiz, 1862 
Rhizocaulus Stechow, \9\9b 
*Silicularia Meyen, 1834 

CLYTIINAE nom. nov. 
Clytia Lamouroux, 1812 
*Gastroblasta Keller, 1883 
*Tulpa Stechow, 192 la 

OBELIINAE Haeckel, 1879 
Gonothyraea Allman, 1864a 
HartlaubellaPoche, 1914 
Laomedea Lamouroux, 1812 
Obelia Peron & Lesueur, 1 8 1 Oa 

p. 50 
p. 51 

p. 58 
p. 67 
Ralph, 1957;Stepanyants, 1979 

p. 69 

p. 71 

Kramp, 1961 

Ralph, 1957;Stepanyants, 1979 

p. 91 
p. 92 
p. 94 
p. 97 
p. 112 

p. 50 
p. 51 

p. 50 

p. 72 
p. 70 

Taxonomy and the phenotype 

The long synonymies and remarks sections in this paper and in that on Obelia (see 
Cornelius, 1975a) reflect the fact that wide phenotypic variation has led to nominal taxa 
being based on unimportant characters. Even the usually reliable Hincks (1868) was misled; 
but in fairness it should be remembered that Hincks, and his colleagues Alder and Allman, 
were breaking new ground when seriously considering the taxonomy of this family. Now, 
with a century of hindsight, the taxa they and others proposed can be better assessed. The 
early workers did not realize how much these hydroids vary, compared with the usually 
much less variable species of Haleciidae, Sertulariidae and Plumulariidae which they knew 

Some of the literature on variation in the species of this and other thecate families has been 
reviewed recently (Cornelius, \915a, b, 1979). The notes in the \915a paper, on Obelia, are 
now summarized and this is followed by some new comments. Authorities for the species 
names are mostly omitted here since they are adequately indicated in the Taxonomic Section 
(P. 47). 


Colony size in erect species probably increases till mechanical breakage occurs (Crowell & 
Wyttenbach, 1957, in Laomedea Jlexuosa; Cornelius, 1975a, in Obelia dichotoma) and is of 
little taxonomic value. Obelia geniculata growth was studied by Ralph (1956) and Ralph & 
Thomson (1968) in New Zealand. Growth was faster in cool conditions than warm. 

Polysiphonic stems occur sometimes in Obelia dichotoma (cf. Millard, 1973) and Laomedea 
neglecta, and are usual in Hartlaubella gelatinosa and Rhizocaulus verticillatus. The species 
Obelia plicata Hincks, 1868, was based on O. dichotoma material of this kind (p. 119). 
Occasional overgrowth of one colony by another of the same species is a rather different 
phenomenon now called auto-epizoism (p. 1 19). 

Branching was most frequent in low water temperatures in Obelia geniculata in New 
Zealand (Ralph & Thompson, 1968). Similar results were obtained from 'Clytia attenuata 1 
by West & Renshaw (1970) who discussed the taxonomic implications (see also notes on 
Growth, p. 42). 

Internode length, extent of annulation, curvature, amount of asymmetric thickening and 
angle of flexure have all be used to define species limits in the Campanulariidae; but only 
asymmetric thickening (in Obelia geniculata} and sometimes curvature (in Laomedea 
Jlexuosa) seem good characters (Cornelius, 1 975a; below, p. 1 1 3). 

Perisarc tanning has been included in some species descriptions, for example in that of the 
now discredited Obelia longissima (discussion in Cornelius, 1975a), but has not been taken 
as a unique species character. The intensity of tanning increases with age in many species. In 
some the perisarc does not seem to darken appreciably, but this might be due simply to its 
thinness. Knight (1965, 1970, 1971) studied the tanning process in Laomedea Jlexuosa. 

Hydranth characters are not useful at species level but the shape of the hypostome is 
consistent throughout some genera. Tentacle number is usually too variable to be useful, 
notably in Clytia hemisphaerica and Obelia dichotoma, but it has been used in defining 
Orthopyxis crenata. 

Hydrothecal characters are useful in this family. But despite their confusing intraspecific 
variation (Broch, 1910, and later workers) we hardly know how the characters develop (e.g. 
Berrill, 1949, in Obelia; Berrill, 1950, in Clytia hemisphaerica, Laomedea Jlexuosa & 
Orthopyxis integra; Knight, 1965, in Laomedea Jlexuosa; Beloussov, 1973, in Gonothyraea 
loveni). Several authors have reported chitinous structures in the hydrothecae of 
Gonothyraea loveni and Obelia bidentata which, however, have proved merely to be 
regularly arranged folds in the delicate hydrothecal walls. Minor variations in the pattern of 
cusps on the hydrothecal rims of several of the species have been given undue weight by 
some authors (see Remarks under O. bidentata). Presence or absence of the hydrothecal 
diaphragm is a subfamily character but is not useful at genus or species level. 

Mammen (1965) noted that an oblique hydrothecal diaphragm viewed from the 'front' 
appears transverse in optical section a point not realized by some taxonomists. It follows 
that rotation of a sloping diaphragm produces a whole series of angles, from horizontal to the 
true maximum slope. Further, even when correctly viewed, a transverse diaphragm is not a 
consistent character (Cornelius, 1 975a). 

Naumov (1969 : 123) stated that many hydroids have larger hydrothecae in cool waters 
than warm, but offered data in only one species (Orthopyxis integra). Although the relation 
might well be valid in many species, detailed proof is needed. Possibly it has already caused 
taxonomic confusion since large, northern specimens of Clytia hemisphaerica have been 
referred to the invalid C. gigantea by several authors (p. 8 1 ) but not all of the large 
specimens were from cool areas. 

Hydrothecal pedicels are variable in length, in amount of annulation and in the presence or 
absence of a smooth central portion. Most species vary widely in these characters. Pedicels of 
reptant species are usually longer than those of upright colonies, perhaps in response to 


greater feeding opportunities away from the substrate than close to it. A peculiar case is the 
long hydrothecal pedicels of floating colonies of the usually reptant Clytia hemisphaerica, 
which have often been regarded a distinct species (C. sarsi nom. nov., p. 78). Implicitly, 
minor variations in pedicel length are phenotypic and overall length ranges are genotypic; 
but evidence is lacking. 

A more useful pedicel character than length is the occurrence of spiral grooving in some 
species in place of the more usual annuli. Spiral grooving seems constant in those species 
having it, and has not been found even exceptionally in annulated species; but annulated 
pedicels have been recorded in the usually smooth to spiral Rhizocaulus verticillatus 
(p. 68). When present a spirally ornamented pedicel is a safe character. 

Another reliable character is the formation in some species of a sub-hydrothecal 
'spherule', first described by Ellis (1755) and noted by many subsequent authors. It is formed 
between two annuli just below the hydrotheca and spaced apart on the pedicel by a distance 
about equal to its width (Fig. 6). Some authors have regarded possession of spherules by 
distinct species as indicating affinity, and this seems usually justified. But spherules are 
simple modifications of existing structures and might have evolved more than once. Thus 
Clytia hummelincki is alone in its genus in having a spherule. 

The spherule structure is otherwise found in the genera Campanularia, Orthopyxis and 
Rhizocaulus. It recalls the basic arthropod joint in having structures analogous to arthrodial 
membranes, but in other ways it resembles the vertebrate ball-and-socket joint. Considerable 
passive flexibility is achieved with little materials, enabling the hydrotheca to be rapidly 
orientated downstream in response to local water movement. It would seem that fewer 
materials are needed in this arrangement than would be needed to construct a rigid, 
unbending pedicel which could maintain the hydrotheca broadside on in strong currents. 
The spherule joints seem to be an evolutionary advance on the simple annulations seen in 
many species of Campanulariidae. Uniformly annulated pedicels bend a little at each 
annulation and have tissue-attachment problems associated with repeated asymmetric 
compression along their whole length. A hydrotheca supported by a spherule can simply 
flip-flop from side to side in response to local current surges, and bends just at one point. In 
addition, spherules would appear more resistant to vertical compression than annuli. 

Murdock (1976) considered very briefly the role of annuli in Obelia sp. main stems. He 
observed that they help bending, an obvious conclusion not often repeated. Hughes (1980) 
studied Laomedea flexuosa and Obelia dichotoma at a few sites on British coasts and found 
higher numbers of pedicel annuli in sheltered situations than exposed. 

Gonothecal shape provides good taxonomic characters in Laomedea but in Clytia, Obelia 
and Orthopyxis it does not. Some nominal species have been based on immature gonothecae, 
for example the invalid 'Laomedea conferta' (p. 104). 

Most species have monomorphic gonothecae but strongly marked sexual dimorphism does 
occur. It has caused taxonomic confusion in Laomedea calceolifera, which was formerly 
regarded as two species. The two kinds of gonothecae proved to be male and female of the 
same species. Gonothecae of L. angulata, L. flexuosa and Gonothyraea loveni tend towards 
sexual dimorphism, and the few L. pseudodichotoma specimens available indicate it occurs 
in that species too. In the other species described herein the gonothecae are monomorphic, 
so far as is known. 

Nematocysts have not yet proved useful in delimiting species in Obelia (Cornelius, 19750) 
but little is known of their potential value in the rest of the family (review in Ostman, 1979). 
As in most hydroids, they are among the smallest of nematocysts and their study requires 
refined techniques. Although Ostman reported slight differences in basal armature between 
the nematocyst threads of some pairs of species here regarded conspecific [Clytia 
hemisphaerica (Linnaeus, 1767) and C. sarsi nom. nov. ( = Laomedea gracilis Sars, 1850); 
Obelia dichotoma (Linnaeus, 1758) and '0. longissima (Pallas, 1766)'], rather few 
populations have yet been studied. She commented that microbasic mastigophores have 


hitherto been separated on characters which are optical artefacts, a conclusion borne out by 
some unpublished electron micrographs she has kindly shown me (Ostman, pers. comm.). If 
the populations having distinct, although very similar, nematocyst types can be 
distinguished on other characters also then Ostman's conclusions will be corroborated. But 
the undischarged capsules are only 6 urn to 8 um long, and the fine details of thread 
structures are unlikely to prove useful in routine identification. Some correlation with gross 
morphological characters would seem desirable. See also page 78 concerning 'Clytia gracilis 
(Sars, 1850)'. 

Habitat preferences and substrate associations are discussed under ecology. Few members of 
the family have a regular habitat association and most seem able to live on a wide variety of 
substrates. The prime exception is Laomedea angulata, living exclusively on eel grasses; but 
since other hydroids live there too this does not help in identification. 

Medusa generation Russell (1953) showed best the extent to which intraspecific variation in 
hydromedusae has led to many invalid species being described; but the factors controlling 
this apparently phenotypic variation are almost unknown. 

The problems surrounding the two nominal species of Obelia recognized from the medusa 
stage in British waters still remain (summarized in Cornelius, 19750, but based largely on 
Russell, 1953 and pers. comm.). In addition there is no clue as to why Obelia medusae on 
release should not always be at the same stage of development. Some authors have based 
nominal species of Obelia partly on tentacle number at the time of release, and although this 
seems unwise the cause of the variation is still unexplained. 

In contrast, four tentacles on release is characteristic of the medusae of most of the Clytia 
species (see generic diagnosis, p. 71). Adult medusae of Clytia linearis, C. hummelincki, C. 
paulensis and Orthopyxis crenata (but see p. 59) are undescribed, as are the hydroid stages 
of C. discoida, C. pentata, C. islandica and arguably C. mccradyi. When all stages in the life 
cycles of these species are known some of the nominal taxa may fall. 

It is remarkable that only a little taxonomic confusion has resulted from the unusual habit 
in Orthopyxis Integra of releasing medusae on some occasions and not on others (p. 63). 

Growth is affected by many factors and has been widely studied in this family. Hammett 
(1943) is now thought to have studied growth in Laomedea Jlexuosa, not Obelia geniculata 
as he stated (Crowell, 1957; Cornelius, 19750). Toth (1969) studied colony 'senescence' in L. 
Jlexuosa. Wyttenbach, Crowell & Suddith (1973) reviewed their own work on stolon 
elongation in thecate and athecate hydroids, treating Laomedea calceolifera, L. Jlexuosa and 
Gonothyraea loveni among the present family. They reported similar results in the two 
Laomedea species but found generic differences in the growth cycles of the stolon tips. 
Cyclic increases in length had been demonstrated earlier in L. Jlexuosa by Wyttenbach 
(1968, 1969) alone; and Beloussov (1961) and Hale (1964) had still earlier found the same 
peculiar phenomenon in stolons of Laomedea Jlexuosa and Clytia hemisphaerica 
respectively. Hale (19730, b) later reported further morphogenetic work on C. hemi- 
sphaerica stolons and reviewed the literature. Beloussov's (1973) important paper described 
more work on the stolons of G. loveni but he did not take environmental factors into account. 
Nevertheless, phenotypic responses to changes in temperature and feeding opportunity are 
known to occur in the stolon of L. Jlexuosa (e.g. Crowell, 1957, 1961), mainly in 'alterations 
largely due to the sensitivity of zones of prospective growth'. These observations are 
interesting in themselves, but their experimental requirements make them unsuitable for 
regular taxonomic use. 

Phenotypic response to temperatue change was noted in the hydroid stage of 'Clytia 
attenuata" by West & Renshaw (1970) who incidentally regarded that species as valid (but 
see p. 40). In vitro colonies at 13-15C were unbranched and could not be distinguished 
from ''Clytia cylindrica Agassiz'; but at 17-19C a kind of branching occurred which these 
authors considered characteristic of C. attenuata. Whatever the validity of the two nominal 
species involved, West & Renshaw drew attention to a taxonomic difficulty resulting from 


phenotypic response. Their extensive review and discussion mainly concerns western North 
Atlantic species ofClytia hydroids and medusae, and further comment would be out of place 

Finally Stebbing (1976, 1979, 1981; see also p. 107) has studied the influence of toxic 
inorganic ions on the growth and death of Laomedea Jlexuosa. He found that growth was 
actually enhanced at sub-inhibitory concentrations of the toxic ions. It seems that care 
should be taken when assessing the morphological characters of material collected from 
slightly polluted places. 


Associations. Only one of the species included here has an apparently obligatory substrate 
association: Laomedea angulata, which has been reliably recorded growing only on eel 
grasses. Obelia geniculata tends to occur on brown algae whereas the very similar O. 
dichotoma grows usually on other substrates; but each occurs occasionally on the substrate 
more usual for the other. The other North Atlantic species of the family show no marked 
substrate associations. They repeat the general hydroid pattern of a few species having some 
substrate specificity with the majority having only loose associations or none at all; but the 
association of L. angulata with eel grasses is unusually close. Nishihira (1968) reported 
'Clytia edwardsia' to be 'characteristic of Zostera marina in Japanese waters. However, 
both Nishihira and Picard (1955, in Algeria) reported many hydroid species growing on 
Zostera but not confined to it. 

Brackish water and estuarine species. Many species of Campanulariidae are tolerant of 
reduced salinity. But while many of the species included here occur either occasionally or 
habitually in low salinity areas, none has been reported from fresh water. The species 
comprise more than a third of the present faunal list: Laomedea angulata, L. calceolifera, L. 
neglecta, Clytia paulensis, Gonothyraea loveni, Hartlaubella gelatinosa, Obelia bidentata, 
O. dichotoma and O. geniculata. All records refer to the hydroid stage and none to the 
medusa. Further details are given in the Habitat sections of these species. 

A similar impression of the family was given by Calder (1976). He found as many as 40 
brackish water hydroids in South Carolina, and of these no fewer than ten were from the 

Interactions between species. Although the phenomenon of overgrowth has been studied in a 
variety of coelenterates and other colonial invertebrates (review in Larwood & Rosen, 1979), 
among hydroids it has been recorded infrequently. Sustained overgrowth of one hydroid 
colony by another is unusual, and in the present family there are a few instances only. For 
example, occasional colonies of Obelia dichotoma with erect stems comprising more than 
one hydrocaulus were once regarded a distinct species (O. plicata, p. 1 19); and the regularly 
polysiphonic stems of some other species treated here may be derived in the same way 
(Rhizocaulus verticillatus, p. 67; Hartlaubella gelatinosa, p. 95; Obelia bidentata, p. 113). 
Overgrowth has been recorded occasionally in Laomedea neglecta (p. 107) but the species is 
not well enough known for this to be assessed. Millard (1973) listed several species of 
thecates from other families showing growth of one colony on another, and introduced the 
descriptive term auto-epizoism. 

Antagonism between colonies is widely known among other coelenterates and in many of 
the invertebrate phyla (Larwood & Rosen), but like overgrowth has seldom been reported 
among hydroids. Hughes (1975) reviewed work on a few species of Campanulariidae ('Clytia 
volubis Packard', C. hemisphaerica, Obelia dichotoma); while in another family Warburton 
(1953) recorded aggression between a colony of Hydractinia echinata (Fleming, 1828) and 
one ofPodocoryne ?carnea Sars, 1 846, on a gastropod shell inhabited by Pagurus sp. 


Geographical distribution 

Most species of Campanulariidae are widely distributed, some occurring nearly throughout 
the World in shallow waters. For example, of the 18 species recorded from New Zealand no 
fewer than seven occur also in British waters (Ralph, 1957). The corresponding figures for 
southern African seas are 21 species and 8 (after Millard, 1975; the immigrant Gonothyraea 
loveni would be additional), and for the antarctic area 13 and five (after Stepanyants, 1979, 
with some taxonomic revision). But few species of the family appear uniformly distributed, 
and many have a patchy local distribution. The local variations are best documented in 
European waters, towards which the following notes on the eastern North Atlantic species 
are unavoidably biased. Further details are given in the Taxonomic Section under each 

Orthopyxis Integra. Although one of the most nearly cosmopolitan of all shallow-water 
hydroids this species has not been found in the Kattegat, Skagerrak, Baltic Sea and Dutch 
waters; and has only occasionally been recorded from Belgium, western Scotland and the 
Irish Sea. 

Clytia hummelincki. So far this species has been reported only from the West Indies, Florida, 
Massachusetts, South Africa and Ghana (p. 83). 

Clytia paulensis. Known for some years from parts of NW France but only recently added to 
the British faunal list (p. 89). However, a specimen collected in S Devon in 1899 has now 
been correctly identified. In the 1970s the species was found in Devon and Suffolk. 

Gonothyraea loveni. In South Africa this species is known from Cape Town docks only, and 
Millard (1975) considered it had spread from Europe to the Southern Hemisphere on ships. 

Hartlaubella gelatinosa. The several nineteenth century Scottish records contrast with a 
single Scottish record this century, in 1932 (p. 95). But the species is still common at least as 
far north as NW England (J. Clare, pers. comm.) and the lack of recent Scottish records may 
be misleading. 

Laomedea angulata. There are few reliable records from the British Isles this century, in 
contrast to an abundance of nineteenth century records (p. 100). Apparently the species has 
yet to regain its former distribution after the temporary decline of the Zostera beds in the 
1930s (described by Tutin, 1942). Although L. angulata was recorded from the Scillies in 
1967 (Robins, 1969) there is apparently no other reliable British record since those from S 
Devon before 1910 (Marine Biological Association, 1957). (But see Addendum.) 

Laomedea calceolifera. Although widespread in North Atlantic waters, relatively con- 
spicuous, and distinctive when fertile, this species has been reliably recorded only twice from 
British waters (S Devon, c. 1 87 1 , by Hincks, 1871; Norfolk, in 1 95 1 , by Hamond, 1957), with 
a third dubious record (Norfolk, in 1899, quoted by Hamond, 1957). Probably the species 
reaches its northern limit in southern England but the paucity of British records is still 
remarkable as the species is well known from NW France (p. 104). 

Laomedea pseudodichotoma. This species has yet to be recorded away from the coastal 
waters of tropical W Africa but it would be remarkable if this indicated the true geographical 
range (p. 1 12). 

Obelia bidentata. This species was first reported from British waters only some 25 years ago; 
and the first record from the south coast of England is reported here (p. 115). It seems 
unlikely that the nineteenth century British collectors would have overlooked so distinctive 
a species, and the absence of earlier records may be genuine. The species was not known in 
Europe until the 1900s. Indeed, it may have been an immigrant into E Atlantic waters from 
the American coast, but this is not certain and early confusion with Hartlaubella gelatinosa 
is not excluded as a reason for the absence of earlier records. See also the next species. 


Obelia dichotoma & O. geniculata. There are records of these widespread species on many 
swimming vertebrates, including a turtle, a shark, the blueback herring and a sea-horse; and 
also on drifting kelp (pp. 117, 118, 1 20). Taken together the records suggest that these species 
might be transported over vast distances. The potential was realized long ago in other 
hydroid families (e.g. Alcock, 1892; Lloyd, 1907; Heath, 1910; review in Gudger, 1928, 
1937). Transport might explain the appearance of the previous species in European waters 
around the 1900s, carried perhaps by ships rather than by vertebrates. However, turtles 
regularly cross the Atlantic (Parker, 1939; E. N. Arnold, pers. comm.) so that natural means 
are not excluded. Clytia hemisphaerica, another widespread species, has similarly been 
found on fish, attached to their crustacean ectoparasites (p. 77). It seems plausible that 
continuous transport across deep ocean basins will promote exchanges between the gene 
pools of these species on different continental shelves; and might explain why many hydroid 
species are virtually cosmopolitan at shelf depths. Paradoxically, it may be that the hydroid 
stages of such species sometimes travel further than their medusae which live for just a few 

Key to species (hydroid stages) 

Many of the species of Campanulariidae are so variable that overlap in characters occurs, 
and identification by a dichotomous key is not always possible. Young and infertile 
specimens are particularly difficult and even with the help of long Museum series some 
specimens cannot be identified. Close study of a single, undamaged hydrotheca is often 
useful and the outline of the unabraded rim can be diagnostic. Hydranth characters are 
seldom useful for identification in this family. 

Provisional identification can frequently be made from the illustrations of the gonothecae. 
Characters based on reproductive structures are mostly omitted from the key, however, as 
many specimens are infertile. When identifying fertile material it can be helpful to determine 
whether the ova develop within the gonotheca or in an external acroyst, and if the gonotheca 
contains developing medusae; but these characters too are mostly avoided in the key. 

1 Colony with erect stems each supporting several to many hydrothecae .... 2 
Colony mainly stolonal, each stem or pedicel supporting one or just a few hydrothecae . 1 8 

2 Rim of hydrotheca even to sinuous 3 

Rim of hydrotheca definitely cusped [Rims often abrade smooth in Gonothyraea loveni, 

Hartlaubella gelatinosa and Laomedea neglecta] 10 

3 Terminal region of hydrotheca flared 4 

Terminal region of hydrotheca not flared 5 

4 Gonothecal aperture narrow; recurved in mature 9 . Laomedea calceolifera (p. 102; Fig. 18) 

- Gonothecal aperture broad, never recurved [tropical] 

Laomedea pseudodichotoma (p. Ill; Fig. 2 1 ) 

5 Internodes curved 6 

Internodes straight 8 

6 Hydrotheca thickened, sometimes much so Obelia geniculata (p. 119) 

Hydrotheca with little or no thickening 7 

7 Hydrotheca usually 1^ times long as broad, or longer; gonothecal aperature usually raised; 

releases medusa Obelia dichotoma (p. 1 1 7) 

- Hydrotheca not much longer than broad; gonothecal aperture not raised; no medusa 

stage Laomedea flexuosa (p. 105; Fig. 19) 

8 With sub-hydrothecal spherule [a locally distributed species] Clytia hummelincki (p. 82; Fig. 10) 
Lacking sub-hydrothecal spherule 9 

9 Internodes rigidly straight; terminal tendrils present in autumn; gonotheca borne on stolon; 

no medusa [on eel grass] Laomedea angulata (p. 98; Fig. 17) 


- Internodes slightly curved; terminal tendrils unusual; gonotheca nearly always axillary; 

medusa released [on many substrates, but including eel grass] . Obelia dichotoma (p. 1 1 7) 

1 Hydrothecal cusps sharp 11 

- Hydrothecal cusps blunt, square, notched or rounded 15 

1 1 Hydrothecal cusps usually bimucronate (hard to see; four species difficult to separate when 

immature) 12 

Hydrothecal cusps all same length 

Clytia hemisphaerica (p. 73; Fig. 9; see also C. mccradyi p. 87, Fig. 13) 

12 Mature colony tall and bushy, with polysiphonic stem and second-order branching; 

gonothecal aperture usually raised, slightly narrower than gonotheca 

Obelia bidentata (p. 1 13) 

Mature colony small, with up to c. 20 hydranths; stem monosiphonic (except occasionally 
in L. neglecta); gonothecal aperture not raised, as broad as gonotheca .... 13 

13 Hydrothecal pedicels longer than hydrotheca . . . . Clytia paulensis (p. 88; Fig. 14) 
Hydrothecal pedicels roughly same length as hydrotheca or shorter 14 

14 Each internode of constant diameter; hydrothecal cusps strengthened by substantial chitinous 

strip; medusa released [Mediterranean southwards] . . Clytia linearis (p. 84; Fig. 12) 

- Internodes slightly bulging; hydrothecal cusps without strengthening strip (folds in 

hydrothecal wall can be confusing); no medusa [Mediterranean northwards] (see also young 
Obelia bidentata) Laomedea neglecta (p. 107; Fig. 20) 

1 5 Hydrothecal margin with rounded cusps 16 

Hydrothecal margin with square cusps, often notched (abrade easily) 17 

16 Mature colony large, polysiphonic; most hydrothecae with subhydrothecal spherule; no 

medusa stage Rhizocaulus verticillatus(p. 67; Fig. 7) 

Mature colony not usually large, always monosiphonic; no spherule; medusa released 

Clytia hemisphaerica (p. 73; Fig. 9) (also C. mccradyi, ?S France only; p. 87; Fig. 13) 

17 Small slender colony, stem monosiphonic; primary branching only; medusa retained as 

gonomedusa external to gonotheca .... Gonothyraea loveni (p. 92; Fig. 15) 

Large bushy colony, stem polysiphonic; with secondary branching; large ova, developing 
into planulae within gonotheca; no medusa. . Hartlaubella gelatinosa (p. 95; Fig. 1 6) 

18 Sub-hydrothecal spherule present 19 

Sub-hydrothecal spherule absent 24 

19 Rim of hydrotheca even 20 

Rim of hydrotheca cusped or undulating 21 

20 Hydrotheca usually much thickenend [common] 

Orthopyxis integra (p. 60; Fig. 6) (also O. crenata with even hydrothecal rims; see text) 

- Hydrotheca unthickened [scarce] Clytia hummelincki(p. 82; Fig. 10) 

21 Hydrotheca > 0-5 mm long, with lines running downwards from rim (sometimes absent in 

one species) 22 

Hydrotheca < 0-5 mm long, without lines 23 

22 Lines meeting tips of cusps, which are roundly pointed, not notched; hydrotheca 2+ times 

long as broad [probably not south of Newfoundland and Spitzbergen] 

Campanularia crenata (p. 52; Fig. 2) 

Lines meeting bottoms of embayments; cusps flat-topped with notch; hydrotheca up to 
1*5 x long as broad [widespread] Campanularia hincksii (p. 53; Fig. 3) 

23 . Hydrotheca usually much thickened [probably Mediterranean southwards] 

Orthopyxis crenata (p. 58; Fig. 5) 

- Hydrotheca unthickened [Mediterranean northwards] . Campanularia volubilis (p. 55; Fig. 4) 

24 Hydrothecal cusps bimucronate [S England southwards] . Clytia paulensis (?. 88; Fig. 14) 

- Hydrothecal cusps simple [widespread] 

Clytia hemisphaerica (p. 73; Fig. 9; also C. mccradyi, ?S France only; p. 87; Fig. 13) 


Taxonomic section 

Family CAMPANULARIIDAE Johnston, 1836 

DIAGNOSIS. Colonial Thecata (sens. Millard, 1975); hydroid stage stoloniferous or erect, 
stolon when present may be branched or unbranched; hydrotheca bell-shaped, radially 
symmetrical, pedicellate, with diaphragm and an associated annular thickening, or with 
annulus alone and without diaphragm; no operculum; hydranths radially symmetrical, 
usually with prominent hypostome; no caecum; one ring of tentacles; nematophores absent; 
d" & 9 gonothecae usually externally identical. Medusa generation when present variable; 
reduced in Obelia and (?) facultatively retained in Orthopyxis; typical leptomedusan in 
Clytia and Gastroblasta; always retained as gonomedusa in Gonothyraea; identifiable in 
more reduced form in at least some other genera and species. 

REMARKS. The family was first proposed by Johnston (1836, 1847, but not 1838). Originally 
Lafoea dumosa (Fleming, 1 820) was included in its scope but was removed to the Lafoeidae 
by Hincks (1868). The limits of the family have remained unchanged ever since and the 
redefinition by Millard (1975) seems sound. Ralph (1957) also provided a detailed appraisal. 

Much has been written about generic limits within the family but it is convenient to go no 
further back in the literature than the works of Broch (1905, 1910) and Goette (1907). Broch 
recognized two broad genera. These he called Campanularia Lamarck, 1816, which had no 
hydrothecal diaphragm, and Laomedea Lamouroux, 1812, which had one. He divided his 
concept of Campanularia into the subgenera Eucampanularia Broch, 1910, having sessile 
gonophores, and Clytia Lamouroux, 1812, with free medusae. Broch split his other broad 
genus concept, Laomedea, into the three subgenera Eulaomedea Broch, 1910, with sessile 
gonophores and no medusoid structures; Gonothyraea Allman, 1864a, with retained 
'eumedusoids' (now called gonomedusoids, p. 93); and Obelia Peron & Lesueur, 1810a, 
with free medusae. Thus Broch's basic division within the family was on a hydrothecal 
character; while within each of the two main divisions his classification was on the state of 
reduction of the medusa generation (following and elaborating on the interpretation of these 
structures by Goette, 1 907). 

Splettstosser (1924 : 424^425) followed Broch's system but further split Broch's subgenus 
Eulaomedea into one group with intracapsular gonophores ('Laomedea gelatinosa\ L. 
flexuosa and L. calceolifera] and a second, in which the mature gonophores were extra- 
capsular (L. neglecta). Splettstosser acknowledged that the classification might be criticized 
since just a small number of species was considered. But Broch (1928) gave it support when 
he later introduced the subgeneric name Paralaomedea for the 'L. neglecta group', 
comprising that species alone, in the combination 'Laomedea (Paralaomedea) brochi 
Splettstosser (= Laomedea neglecta Alder)'. [Splettstosser's restriction of the subgenus 
concept was thus cited as indication by Broch; but the authority for the associated subgenus 
name Paralaomedea was Broch (1928).] Finally Hummelinck (1936) redefined the subgenus 
rather tightly, again to include only L. neglecta. 

As Splettstosser had commented, very few species were considered in his classification and 
it is questionable whether so many infra-generic divisions were justified. (Some additional 
sub-divisions of Eulaomedea' proposed by Splettstosser were not given names, and are not 
mentioned here.) His own work on L. neglecta and other species, and that for example of 
Goette (1907) and Miller (1973), emphasized that the gonophore 'types' identified by 
Splettstosser form part of a series in which the medusa is progressively reduced. The 
gonophore of L. neglecta seems simply to fit into this series. Further, Broch's (1910) primary 
division of the family into two was on the basis of a single hydrothecal character, and this 
division too might be challenged. 

The extent to which the medusa-medusoid-gonomedusoid-gonophore series should be 
classified into genera will perhaps be debated for as long as the series is regarded valid. But 
today as in Broch's time, most is known about the life-cycles of the western European 


species. Until more information is available on species from other parts of the World it seems 
unwise to split Laomedea into subgenera. 

Aside from the taxonomic debate, there are some nomenclatural problems which need 
solution. These I have considered in a submission to the International Commission on 
Zoological Nomenclature (Cornelius, 1981). The aims of the proposals are provisionally 
included in the present paper. 

The subgenera proposed by Broch, Splettstosser and others need not be recognized. But 
the evolutionary fate of the medusa generation is still reflected in the classification adopted 
here. In Orthopyxis the medusa is reduced and lacks several normal adult characters, 
functioning simply as an ephemeral gamete carrier. It is thought to be facultatively released 
in some or all of the Orthopyxis species (see p. 63); while in the closely related Orthonia it is 
still further reduced, to a retained acrocyst. In Campanularia, Rhizocaulus and Silicularia 
the gonosome has become intracapsular, with no obvious indications of a medusoid 

A parallel series showing progressive retention of the medusa can be demonstrated within 
the genera Obelia, Gonothyraea, Laomedea and Hartlaubella. In Obelia the medusa is 
released. The extra-capsular gonomedusoids of Gonothyraea are easily identified as retained 
and vestigial medusae; and in Laomedea it has been shown that the gonosomes of several 
species represent reduced medusae. Indeed, so reduced has the medusa generation of 
Laomedea become that until the work of Miller (1973; see also Goette, 1907) the medusoid 
nature of the gonophore was not appreciated. The same confusion prevailed also in 
interpretations of the reproductive structures in Orthopyxis Integra, in which the medusa is 
sometimes retained. It was thought until quite recently that the retained examples had 
'sporosacs' in place of medusae, and that they might therefore be a different species (O. 
caliculata', p. 65-66)! 

The genera Clytia, Gastroblasta and Tulpa, which have a true hydrothecal diaphragm and 
sub-hydrothecal spherules, apparently form another group but their relation to the rest of the 
family is not clear. 

The three series recognized are shown in Figure 1 . The groupings seem natural and are 
here given subfamily status: Campanulariinae (p. 50), Clytiinae nom. nov. (p. 69) and 
Obeliinae Haeckel, 1 879 (p. 9 1 ). See also page 49. 

The generic limits suggested by Millard (1975) are slightly modified, as is her 
nomenclature. Orthopyxis is here separated from Campanularia sens. Millard. Eulaomedea 
sens. Millard is here called Laomedea. Sertularia gelatinosa Pallas, 1766, not in Millard's 
faunal area, is referred to the monotypic genus Hartlaubella; and another species not in her 
list, S. verticillata Linnaeus, 1758, is here referred to the nearly monotypic Rhizocaulus. 

To promote stability of nomenclature I have attempted to include all extra-limital generic 
synonyms. The valid genera found outside the NE Atlantic are treated briefly. They are 
Eucalix, Orthonia and Silicularia from the Campanulariinae (p. 50); and Gastroblasta and 
Tulpa from the Clytiinae (p. 70). The problem genus Hypanthea is discussed along with 
Silicularia (p. 50). 

From the medusa stage, Kramp (1961) recognized only five genera World-wide: Agastra, 
Eucopella, Gastroblasta, Obelia and Phialidium\ but of these only Gastroblasta and Obelia 
can now be recognized. Reference to discussions of these genera can be made using the index.. . 

I have previously commented (Cornelius, 1975a) on the genera Medusa Linnaeus, 1758 
(part); Schizocladium Allman, 1871; Obelaria, Obeletta & Obelissa, all Haeckel, 1879 
(Obelaria Hartlaub, 1897, is a junior homonym and is discussed here under Hautlaubelld)', 
and Monosklera von Lendenfeld, 1885; all except the first of which fall into the synonymy of 
Obelia Peron & Lesueur, 1810#. I then mentioned also Thaumantias Eschscholtz, 1829, a 
junior subjective synonym of Clytia Lamouroux, 1812 (see below, p. 71). I overlooked 
(p. 254) that Mayer (1910: 262) had designated Sertularia volubilis sens. Ellis & Solander, 
1786 (non Linnaeus, 1758) type species of Clytia (see p. 70 below). Lastly, I have reversed 
my opinion on the use of the genus name Laomedea. 



Medusa entirely 

Medusa reduced 
to external 

Medusa vestigial 
and retained 

Medusa released 

Laomedea (part) 





Campanularia, Tulpa 








Fig. 1 Affinities within the Campanulariidae. The genus Eucalix is not included since its method 
of reproduction is unknown, but vegetative characters suggest it is close to Orthopyxis 
(discussion on pp. 50-51). The diagram shows present-day similarities, not phylogenies, but the 
general direction of evolutionary advance is up the pagejand towards the right. Knowledge of the 
group is incomplete and the chart should be regarded as provisional. 

Some of the species described herein are known only from either hydroid or medusa stages, 
and others were formerly so. Most species in which the two stages have been reconciled are 
now known by appropriate combinations, based on application of the International Code of 
Zoological Nomenclature; but those with incompletely known life-cycles cannot yet have 
their names confidently derived. Current knowledge in this family seems adequate for the 
Code to be applied to the two stages simultaneously, but this is arguably not so in all hydro- 
medusan families and in some there may still be a case for retaining the dual system. 

The subfamily divisions and their nomenclature 

Although the limits of the family Campanulariidae have been agreed for nearly a century 
and a half (p. 47) only three authors (Haeckel, 1879; Mayer, 1910; Russell, 1953) have 
sought to group the genera into formal subfamilies. Indeed, until some quite recent studies of 
the range of reproductive structures found within the family had appeared (Miller, 1973; but 
also Splettstosser, 1 924) interpretation and grouping had been difficult. Miller's important 
work showed that the structures which had once been called fixed gonophores in for example 
Laomedea spp. were vestigial, retained medusae; and that the curious externally-held 
'meconidium' ofGonothyraea loveni is similarly to be regarded as a retained medusa. 

Happily, this new interpretation of the dispersive generations (planulae and medusae) 
corroborates the broad divisions of the family based long ago solely on the vegetative 
characters of the hydroid stage (Broch, 1905, 1910; Goette, 1907; see p. 47, above). Hence 
the subfamily divisions adopted here, which draw on both groups of characters, might seem 
soundly based. But some problems remain and further refinement will no doubt be achieved 
when more is known of the non-European members of the family. 



Campanulariadae Johnston, 1836 : 107 (part). 

Obelidae Haeckel, 1879 : 163 (part). 

Obelinae: Mayer, 1910 : 231 (part). 

Orthopyxinae Russell, 1953 : 303. 

non Campanularinae: Russell, 1953 : 284 ( = Clytiinae nom. nov.; see p. 69). 

NOMENCLATURE. The spelling Campanulariinae takes as its root the genus name 
Campanularia, and Campanularinae is wrong. 

DIAGNOSIS. Campanulariidae with colony usually replant, secondarily erect and poly- 
siphonic in Rhizocaulus; no true hydrothecal diaphragm; medusa absent except in 
Orthopyxis, where reduced. 

TYPE GENUS. Campanularia Lamarck, 1816, the nominate genus. 

SCOPE. The genera Campanularia Lamarck, 1816; Silicularia Meyen, 1834; Orthopyxis 
Agassiz, 1862; Rhizocaulus Stechow, 19196; Orthonia Stechow, 19230; and probably 
Eucalix Stechow, 192 la. 

REMARKS. Of the included genera only Campanularia, Orthopyxis and Rhizocaulus are fully 
treated in this paper. The others have not been recorded from the eastern North Atlantic and 
are discussed only in this section. 

Millard (1975:201) united Campanularia and Orthopyxis because she had seen 
Orthopyxis colonies having some unthickened hydrothecae; but I feel the remaining 
characters justify a separation. 

The genus Silicularia Meyen, 1834, was proposed to include two species, S. rosea and S. 
gracilis, both being described as new. The early date of Silicularia and inadequacies in the 
descriptions of the two species make detailed comments necessary. The type species of 
Silicularia is S. rosea, designated by Stepanyants (1979 : 33). The species was based on 
Ethiopian and South African material. It was redescribed by Nutting (19 1 5) and Stepanyants 
(1979), and Blanco (19670) provided useful notes. The second species, S. gracilis, was based 
on infertile hydroid material from the Sargasso Sea and the Azores. It was probably a Clytia 
species. However, the figures and description do not include details of the hydrothecal rim or 
reproductive structures and I agree with Bedot (1905 : 171) that the species cannot be 
confidently assigned (see also p. 118). Nutting (1915 : 66) referred 'S. gracilis' to the rather 
dubious species Orthopyxis clytioides (Lamouroux, in Freycinet, 1824, as Tubularia). He 
wrongly quoted Meyen as using the combination Silicularia clytioides. Meyen actually used 
S. gracilis. Rees & White (1966) made the same error when citing Meyen's Azores record. 
There seem no other reports of 'S. gracilis' from the eastern North Atlantic. I provisionally 
refer T. clytioides Lamouroux to Obelia dichotoma, under which it is discussed further 
(p. 1 1 8). S. gracilis was mentioned recently by Stepanyants ( 1 979), as Campanularia. 

Nutting (1915), Bedot (1925), Broch (1929) and Stepanyants (1979) all regarded as 
congeneric with Silicularia the later genus Hypanthea Allman, 18760 (type species H. 
repens Allman, 18760, by monotypy; type locality of the type species, Kerguelen I.), and I 
agree. Hypanthia Nutting, 1915 : 22, was a lapsus. Both Allman's (18760, 1888) concept of 
Hypanthea and Nutting's and Stepanyants' of Silicularia included thick, asymmetrical 
hydrothecae, pedicels and stolons reminiscent of Orthopyxis Agassiz, 1862, to which the 
original concepts of the two genera come close. I have not located the type material of H. 
repens, but later material referred to Hypanthea species by Allman (1888) had an 
anastomosing stolon and other orthopyxine features. 

So far as I can determine no Silicularia species has been recorded from the eastern North 
Atlantic. S. atlantica (Marktanner-Turneretscher, 1890, as Hypanthea), was based on 
material said to have come from 6 S, 38 W, but this position is on the mainland of South 

The genus Eucalix Stechow, 19210 : 254, was proposed to accommodate the sole species 


Campanularia retroflexa Allman, 1888, type locality Honolulu. Stechow maintained that 
the unusual hydrotheca of E. retroflexus justified generic separation. Both the type series 
(BMNH reg. no. 1888.11.13.14) and the original illustration (Allman, 1888 : pi. 11, figs 1, 
la) show the hydrothecal characters regarded important by Stechow. These characters, 
together with the anastomosing stolon of the type material, suggest that Stechow was justified 
in proposing the new genus. Millard (1957 : 196; 1975 : 212, as Campanularia morgansi) 
listed relevant literature. (I am grateful to Professor W. Vervoort for discussing the characters 
of this species; and to Dr D. M. Devaney for identifying the substrate of the type material. The 
substrate is a coralline alga, Halimeda sp., not a millepore as Allman stated. Dr Devaney 
informs me millepores have not been recorded from Hawaii.) 

The genus Orthonia Stechow, 19230 : 94, 107, was proposed to accommodate a single 
orthopyxine species, Campanularia everta Clarke, 1876:253-254, pi. 39, fig. 4, type 
locality San Diego. Nutting had subsequently assigned to this species material having 
acrocysts and it was this character on which Stechow distinguished the genus from 
Orthopyxis. An element of subjectivity was thereby introduced since Stechow assumed that 
Nutting had material of the same species as had Clarke; and Stechow had no proof. Further 
revision of the orthopyxine species seems necessary before Orthonia is evaluated further. 

Genus CAMPANULARIA Lamarck, 1816 

Campanularia Lamarck, 1816:112 (part); Hincks, 1868 : 160 (part); Nutting, 1915 : 27 (part). 
Campanula Westendorp, 1843 : 23 (lapsus pro Campanularia}. 
Campanulata Agassiz, 1862 : 354 (lapsus pro Campanularia). 
Campanularia (Eucampanularid) Broch, 1910 : 184 (part). 
Paracalix Stechow, 1923c : 3. 

TYPE SPECIES. Provisionally to be taken as Sertularia volubilis Linnaeus, 1758 : 811 (non 
Ellis & Solander, 1786, see p. 70) as designated * by Naumov (1960: 249). Nutting 
(1 9 1 5 : 28) earlier designated S. verticillata Linnaeus, 1758 : 8 1 1 , as type species but applica- 
tion has been made to the International Commission on Zoological Nomenclature for this 
designation to be set aside (Cornelius, 1981; see Remarks). Broch (1905 : 10) proposed that 
'Campanularia calyculata Hincks, 1853' should be type species, but 'calyculata' was not 
among the species originally included in the genus and so is not eligible. The correct spelling 
is of course caliculata (p. 65). 

DIAGNOSIS. Stoloniferous and colonial Campanulariidae, stolon not anastomosing; hydro- 
thecae borne on pedicels inserted on the stolon at irregular intervals; true diaphragm absent; 
sub-hydrothecal spherule present; no medusa stage. 

REMARKS. The species Sertularia verticillata Linnaeus, 1758, was designated type species of 
Campanularia by Nutting (1915). But some authors, with whom I agree, have sought to 
remove verticillata to a distinct genus (Stechow, 19 \9b, c; Naumov, 1960, 1969). This would 
leave the name Paracalix Stechow, 1923c, available for the present genus; so that Paracalix 
would become applied for example to the common hydroids widely known as Campanu- 
laria hincksii (p. 53) and C. volubilis (auct.; p. 55). The genus Paracalix Stechow, 1923c, 
was proposed to accommodate only Campanularia pulcratheca Mulder & Trebilcock, 
1914: 11, pi. 2, figs 1-2, a species based on sterile material from Torquay, Victoria, 
Australia. (The generic name was actually misprinted Cmpanularia in Mulder & 
Trebilcock's heading.) The hydrotheca was sigmoid in lateral view and Stechow was 
impressed by the resulting bilateral symmetry. This was the main character on which the 
species, and subsequently Stechow's proposed genus, were based; but the specimen seems 
simply to have been a deformed specimen of C. volubilis or a closely related species. On this 
interpretation the species pulcratheca and the genus Paracalix are referred to Campanularia. 
I have applied to the International Commission on Zoological Nomenclature for Nutting's 
designation of S. verticillata as type species of Campanularia to be set aside (Cornelius, 
1981). If approved, this will validate Naumov's (1960) designation of S. volubilis Linnaeus, 


1758, as type species of Campanularia. As a consequence the genus name Rhizocaulus 
Stechow, 19196, can then be applied to the species verticillata (in the combination R. 
verticillatus, p. 67). See also the notes under Rhizocaulus (p. 67). 

The subgenus Eucampanularia Broch, 1910, was introduced to embrace the five species 
Sertularia volubilis Linnaeus, 1758, Campanularia Integra Macgillivray, 1842, C. groen- 
landica Levinsen, 1893, C. speciosa Clarke, 1877 and S. verticillata Linnaeus, 1758. The 
subgenus name has hardly been used in the literature. I designate S. volubilis Linnaeus, 1758, 
as its type species; so that Eucampanularia can be regarded a junior objective synonym of 
Campanularia (subject to my proposals to the ICZN being accepted; see also Cornelius, 

Campanularia crenata Allman, 1 8766 
(Fig. 2) 

Campanularia crenata Allman, 1 8766 : 258-259, pi. 11, figs 1-2. 

Campanularia speciosus Clarke, 1877:210 (lapsus pro speciosa). 

Campanularia speciosa (Clarke, 1877:214-215, pi. 9, fig. 11; Linko, 1911 : 185-187, fig. 34 (syn. 
C. crenata Allman); Broch, 1912a : 17-18, fig. 3; Nutting, 1915 : 48, pi. 8, fig. 5 (syn. C. crenata 
Allman); Broch, 1918 : 158-159 (syn. C. magnifica Eraser); Calder, 1970 : 1519, pi. 4, fig. 3. 

Campanularia magnifica Eraser, 1913 : 164, pi. 1 1, figs 1-3. 

NOMENCLATURE. The widely used trivial name speciosa was introduced in a paper published 
on 2 January, 1877, and not in 1876 as usually assumed. Hence crenata, genuinely 
introduced in 1 876, has priority (see note on page 1 29 under Clarke, 1877). 

The combination Campanularia crenata has been applied also to the species here called 
Orthopyxis crenata; but in that species the original binominal was Eucopella crenata, and 
primary homonymy has not occurred. There is secondary homonymy, however, and this is 
discussed under O. crenata (p. 60). 

TYPE LOCALITY AND MATERIAL. Infertile colony on 'Thuiaria crassicaulis* (Sertulariidae), 
?Tsuger Straits, Japan, 183m (lOOfms); 1877.4.12.8 (previously unpublished data with 

OTHER MATERIAL EXAMINED. All BMNH material is listed. 45 m, Store Hellefiskebanke, 
Greenland, fertile colony on Sertularia mirabilis (Verrill, 1873), coll. G. M. R. Levinsen, 
exch. Copenhagen Mus.; 1896.8.15.2 (Fig. 2; ?mentioned, Broch, 1918). 'Greenland', 
infertile colony on Sertularia mirabilis, ex D'Arcy Thompson colln, pres. Univ. Dundee; 
1957.1.1.12. Infertile colony on Sertularella sp., Norman St, Labrador, Canada, ex D'Arcy 
Thompson colln, pres. Univ. Dundee; 1956.10.23.69. Infertile colony on Symplectoscyphus 
sp., Bell I, Newfoundland, Canada, 17 Apr 1892, ex D'Arcy Thompson colln, pres. Univ. 
Dundee; 1957.1.3.24. Infertile fragment, Bel Sund, Spitzbergen, 14 Jul 1898, 20m, coll. 
Spetsberg Expedn, exch. Stockholm Mus.; 1960.8.29.33. No locality, fertile colony on 
sertulariid hydroid, exch. Copenhagen Mus.; 1 9 1 2. 1 2.2 1 .44. 

DESCRIPTION. Colony reptant. Stolon tortuous, branched, rugose. Hydrothecae on usually 
long pedicels, at irregular intervals; large, narrowest c. \ from rim, bulging out basally; rim 
much flared, with c. 10 rounded cusps, usually with striations running proximally from apex 
of each cusp; sub-hydrothecal spherule present; pedicel usually longer than hydrotheca, up 
to c. 3x length, spirally grooved throughout. Gonotheca ?rf = 9, elongate-ovoid, with or 
without long neck, borne on stolon. 

Variation. The short necked gonotheca illustrated was apparently mature, indicating that 
the long necks usually regarded distinctive are not invariably present. 

DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Present material 
(1896. 8. 15. 2) has just one in each gonotheca but there may have been more in life. 



DISTRIBUTION. A circumpolar Arctic Ocean and cold water species recorded in the Atlantic 
as far south as Newfoundland (present material), W Greenland and Spitzbergen (Broch, 
19120;Calder, 1970). 

HABITAT. Usually reported epizoic on other hydroids. Naumov (1960, 1969) found an 
extreme depth range of 3 m to 600 m in Russian seas, most of his records being between 20 m 
and 200 m. 

REMARKS. This species has been widely known by the combination Campanularia speciosa. 

The long gonothecal neck often thought characteristic of the species is not present in all 
the material listed here, and is evidently variable in length. 

Broch's (1918) attack on the validity of Campanularia magnifica Eraser, 1913, left no 
doubt that it is conspecific. 

Campanularia hincksii Alder, 1856a 
(Fig. 3) 

Campanularia hincksii Alder, 1856a:360, pi. 13, fig. 9; Hincks, 1868: 162-163, pi. 24, fig. 3; 
Goette, 1907 : 189-193, pi. 15, figs 307-312; Broch, 1933 : 87-93 (syn. C. aha Stechow); Vervoort, 
\946a : 276-277, fig. 122 (syn. C. aha Stechow); Patriti, 1970 : 33-34, fig. 41 (syn. C. brachycaulis 
Stechow, 1919a, here referred to Clytia hemisphaerica, see p. 82; C. macrotheca Leloup); 
Millard, 1975 : 208, fig. 67b-e. 

Campanularia aha Stechow, 1919a : 54-57, fig. P. 

Campanularia rara Stechow, 1 9 1 9a : 60-6 1 , fig. R. 

Campanularia macrotheca Leloup, 1930a : 101-102, figs 1-3. 

TYPE LOCALITY AND MATERIAL. Coast of Northumberland, England (Alder, 1856a; Millard, 
1975). The syntype series is preserved jointly in the Hancock Museum, Newcastle upon 
Tyne, Northumberland (several colonies in spirit, epizoic on sertulariid hydroids) and the 
BMNH [small dry colony, 1857.8.3.58, epizoic on Lafoea dumosa (Fleming, 1820)]. It has 
been catalogued by Cornelius & Garfath (1980). 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia aha Stechow, 19190, infertile 
fragment of syntype on microslide, Naples; Munich Zoological Museum. 
C. rara Stechow, \9\9a, infertile fragment on microslide, Marseille; MZM. 

OTHER MATERIAL EXAMINED. BMNH collection, c. 50 specimens. The following, collected by 
W. J. Rees, had fertile d" gonothecae: Hjeltefjord, nr Bergen, Norway, 40-90 m, 9 Apr 1962, 
1962. 10.7.20; I of Cumbrae,W Scotland, 90m, 11 Jul 1966; 1967.12.1. 10-12. 

DESCRIPTION. Colony a tortuous stolon bearing unbranched hydrothecal pedicels at irregular 
intervals. Hydrotheca large, campanulate, truncate basally; length : breadth ratio variable 
(1-3-2-25:1, Millard, 1975); rim castellate, 8-15 blunt cusps each usually notched, 
occasionally deeply; main embayments deep, curved, often conspicuous, with characteristic 
folds trailing down from centres. Hydrothecal pedicel long, with spherule distally; shaft 
smooth to sinuous, usually with several annuli basally, sometimes also 1 - several annuli 
along length (Vervoort, 1946a). Hydranth ?undescribed, 18-24 tentacles visible in 
contracted BMNH material. Gonothecae cf = 9, borne on stolon; sub-cylindrical, sometimes 
asymmetrical; broadest near base, truncate below, tapering gradually above; sides smooth to 
irregularly sinuous in a loose succession of rings; truncated and sometimes slightly flared 
distally; aperture wide, terminal; planula development probably internal; gonothecal pedicel 
short, ringed; colonies dioecious. 

Variation. The BMNH series shows variation in the following features: size and 
length : breadth ratio of hydrotheca, height and number of cusps, depth of notch in cusps, 
presence or absence of longitudinal folds in hydrothecal wall; sinuosity of perisarc of 
hydrothecal pedicel, length of pedicel, number of basal annulations (may be absent), shape of 
proximal cavity in hydrotheca; sinuosity of gonothecal wall, amount of flaring below 
gonothecal aperture. Billard (1934) reported that the notch in the tips of the hydrothecal 


cusps may be absent, when the hydrothecae sometimes resemble those of Campanularia 
volubilis(p. 55). 

DISPERSIVE STAGE. Planulae, which probably develop within the female gonotheca. Develop- 
ment of the male gonomedusoid was described by Goette (1907). 

REPRODUCTIVE SEASON. Jun-Oct in NW France (Teissier, 1965). BMNH fertile material has 
collection dates within these limits except a male specimen from near Bergen, dated 9 Apr 

DISTRIBUTION. Nearly cosmopolitan in shallow waters. Although not the most abundant 
hydroid C. hincksii can be expected almost throughout the eastern North Atlantic, local 
conditions permitting. Notable records include: N & S Iceland, Lofoten Is & Norway 
(Kramp, 1938); Mediterranean (Picard, 19586); Italy (Rossi, 1971); Portugal (Da Cunha, 
1950); Cap Spartel, Tangier & Cap Blanc, Morocco (Billard, 1907); Azores (Rees & White, 
1966); Mauritania (Billard, 193 la); South Africa ('rare', Millard, 1975). The species is 
widespread in parts of temperate western Europe, including the British Isles (Hincks, 1868), 
but is scarce in Dutch and Belgian waters (Vervoort, 19460; Leloup, 1952). There are several 
records from the Skagerrak and Kattegat (Kramp, 1935) and W Sweden (Jagerskiold, 1971), 
but no records from the Baltic Sea (Stechow, 1927; Broch, 1928; Naumov, 1960, 1969) or 
Black Sea (Naumov). 

HABITAT. Usually recorded between 20 m and 200 m but occasionally deeper: 'a few metres 
down to 800m' (Kramp, 1938); 20-100 m, SW England (Marine Biological Association, 
1957); c. 20 m, SW Wales (Crothers, 1966); 25-50 m, Scilly Is (Robins, 1969); below 20 m, 
NW France (Teissier, 1965); 1 12-120 m, Strait of Gibraltar & Morocco (Billard, 1907); 
27-98 m, Azores (Rees & White, 1966); 86-210 m, southern Africa (Millard, 1975). Shallow 
records include: 15 m, NW Wales (Knight-Jones & Jones, 1956); 10-1 12 m, Faeroes (Kramp, 
1929); 9-5-80 m, W Sweden (Jagerskiold, 1971). Apparently no intertidal records. 
The species seems unrecorded from brackish waters and may be stenohaline. 

REMARKS. Millard (1975) summarized the doubts concerning the shape of the d gonotheca, 
which it seems has not been reported before now. The BMNH series includes several 
colonies in which the gonothecal contents are preserved. The d 1 and 9 gonothecae are 
identical, and are borne on separate colonies. The contents of the cf were described by Goette 
(1907) and are clearly gonomedusoid in Miller's (1973) terminology. 

The name applied to the distinct but closely related nominal species Campanularia laevis 
Hartlaub (1905:565-567, pi. 1, based on Chile material) is a junior homonym of 
Campanularia laevis Couch, 1844 (see p. 65). I propose the name Campanularia agas 
nom. nov. for the Hartlaub species. C. agas was recently redescribed by Vervoort 
(1972 : 85-87, as Campanularia laevis). Both Hartlaub and Vervoort discussed similarities 
between C. agas (= C. laevis Hartlaub) and C. hincksii. 

Hickson & Gravely (1907) referred additional material to 'C. laevis Hartlaub', but Totton 
(1930) considered their material distinct. He referred it to a third nominal species, 
Campanularia hicksoni Totton, 1930. This was a species proposed to accommodate the 
material described by Hickson & Gravely, and also some collected by the Terra Nova'. It 
was discussed briefly by Rees & Thursfield (1965:90, as Campanularia laevis sensu 
Hickson & Gravely) and in detail by Stepanyants (1979 : 29). 

Campanularia aha Stechow, 19190, was based partly on new material from Villefranche 
and partly on some accounts of earlier authors. Stechow's material had young male 
gonothecae characteristic of C. hincksii, but he illustrated a hydrotheca more typical of 
Clytia hemisphaerica. However, the earlier descriptions included (i.e. those of Billard, 1907; 
Goette, 1907; Broch, 19126) seem undisputedly of C. hincksii. Hence I concur with Broch 
(1933) and Vervoort (19460) in regarding C. aha conspecific, and not with Picard (19510, 
1955) who maintained it distinct. 

Campanularia macrotheca Leloup, 19300, based on material from Monaco, was 
justifiably referred to the present species by Patriti (1970). 



Figs 2-4 Fig. 2 Campanularia crenata. (a) hydrotheca and (b) 9 gonotheca, Greenland, 
1896.8.15.2. Scale 500 //m. Fig. 3 Campanularia hincksii. (a) hydrothecal pedicel and part of 
stolon. The pedicel is unusually short for the species. Note the irregular thickening. W Norway, 
30-40 m. (b) 9 and (c) d gonothecae, sexes identified from contents. W Scotland, 90 m, July 1 966; 
1967.12.1.16 and 10 respectively. The 9 gonotheca is unusually long. Scale (a-c) 500 //m. 
Fig. 4 Campanularia volubilis. (a) hydrotheca and pedicel, 30^0 m, nr Bergen, 1 5 August 1962; 
1962. 1 1 .7.6. Scale 500 um. (b) vertical optical section through (a), showing flexible region. Scale 
50 urn. (c-d) two gonothecae, one with ova, from a single colony, Shetland; 1912.12.21.55. Scale 
as (a). 

Campanularia volubilis (Linnaeus, 1758) 
(Fig. 4) 

Corallina minima scandens, vesiculas campaniformes in summo caule lineari contorto gerens. Ellis, 

1755: 24-25, pi. 14, figs A, a. 
Sertularia volubilis Linnaeus, 1758:811; Linnaeus, 1767:1311; (non Pallas, 1766 : 122, junior 

homonym, = Calycella syringa (Linnaeus, 1767), see also Cornelius, 1978; non Ellis & Solander, 

1 786 : 5 1 , pi. 4, figs E, e, F, f, = Clytia hemisphaerica, see p. 70). 
Sertularia uniflora Pallas, 1766 : 121-122 (nom. nov. pro S. volubilis Linnaeus, 1758; see pp. 77-78); 

(non Ellis, 1768 : 434, pi. 19, fig. 9, = Clytia hemisphaerica, see p. 78). 
Campanularia volubilis: Alder, 1857: 125-126, pi. 4, fig. 7; Hincks, 1868: 160-162, pi. 24, fig. 2 

(non Hincks, 1 852, nee Du Plessis, 1871, = Clytia hemisphaerica, see p. 70). 
Campanularia groenlandica Levinsen, 1893: 168, pi. 5, figs 10-12; Naumov, 1960:252-253, fig. 

139; Naumov, 1969 : 273-274, fig. 139; see Remarks, 
non Clytia volubilis: Hargitt, 1909 : 373-374 ( = C. hemisphaerica, see p. 78). 
Clytia mollis Stechow, 1919a : 44^45, fig. L (?syn. Clytia iaevis Weismann, 1883). 
Campanularia brachycaulis Stechow, 1919a : 62-63, fig. T. 

NOMENCLATURE. Further synonymies were given by Bedot (1901-1925), Vervoort (\946a) 
and Naumov ( 1 960, 1 969) among others. 

TYPE MATERIAL AND LOCALITY. Linnaeus (1758) gave only Ellis' (1755) illustration as 
indication. As with some other hydroids (Cornelius, 1979:309, notes 11-14) Linnaeus 
apparently based the designation on Ellis' plate and not on specimens. Almost certainly the 
material now in the Linnaeus collection in the Linnean Society of London (Savage, 
1945 : 206) reached Linnaeus after the original description was published and cannot be 


regarded as type (Cornelius, 19750 : 273, footnote). The material collected and described by 
Ellis [infertile colony on Hydrallmaniafalcata (Linnaeus, 1758); Brighton, Sussex, England, 
June, 1754; illustrated, Ellis, 1755 : pi. 14, figs A, a] can thus be considered type. Although 
some hydroid material of John Ellis survived until recently it seems that only a single speci- 
men (of Nemertesia sp.) escaped destruction during World War II (Cornelius, \915a : 267, 
footnote) and the specimen illustrated by Ellis can be assumed lost. The type locality is 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia groenlandica Levinsen, 1893, 
infertile syntype material on two pieces of Lafoea dumosa (Fleming, 1 820), in spirit, exch. 
Copenhagen Mus., Davis Strait, '80 frns'; 1 896.8. 1 5.1. 

Campanularia brachycaulis Stechow, 1919a, infertile fragments on 2 microslides, 
Villefranche; Munich Zoological Mus. 

Clytia mollis Stechow, 19 19a, small fertile colony on weed, Sete, S France; MZM. 

OTHER MATERIAL EXAMINED. BMNH collection, c. 60 specimens. 

DESCRIPTION. Colony comprising creeping stolon bearing irregularly spaced, erect, straight 
pedicels each supporting a hydrotheca. Stolon smooth to irregularly spirally grooved. 
Pedicels apparently always unbranched, smooth to spirally grooved throughout; sub- 
hydrothecal spherule present. Hydrotheca tubular, tapering abruptly basally; rim with 10-12 
shallow blunt cusps; sometimes with fine longitudinal striae associated with the cusps. 
Gonothecae scarce, <S = 9, on short pedicels; flask-shaped, smooth, aperture at end of neck of 
indefinite length; neck forms after body of gonotheca; on stolon or (Hincks, 1868) on 
hydrothecal pedicels. 

DISPERSIVE STAGE. Planulae, brooded in the 9 gonotheca. There is no medusa stage. 

REPRODUCTIVE SEASON. Apparently the only published information is of a fertile specimen 
off Norfolk, 16 June, 1951 (Hamond, 1957). None of the dated specimens in the BMNH is 
fertile. Possibly reproduction in this species is usually vegetative. Hamond found fertile 
material just once, and only a few of the BMNH specimens have gonothecae. 

DISTRIBUTION. Common from southern England northwards, but probably present in 
scattered localities further south. Not recorded from NW France or Belgium, nor reliably 
from the Netherlands (Teissier, 1965; Leloup, 1952; Vervoort, 1946a). However, the species 
is well known from the south coast of England (Ellis, 1755; Marine Biological Association, 
1957). Vervoort (1949) recorded a single specimen from the Channel Isles but was 'unable to 
trace records along the NW coast of France'. There are some records from the Mediterranean 
Sea (Stechow, 1923a; Riedl, 1959; Naumov, 1969) but Picard (19586) excluded the species 
from his faunal list. There is a record from N Spain (Santander; Rioja y Martin, 1906) and 
another from Mauritania (Billard, 193 la); but in general there are few records further south 
than the British Isles. 

Northerly records include: Iceland, numerous examples (Kramp, 1938); Greenland, 
widespread up to 72 N (Kramp, 1943); N Norway and N coast of Russia (Mathiesen, 1928; 
and Naumov, 1969, as C. groenlandica). 

HABITAT. Both Mathiesen (1928, N Norway) and Naumov (1969, Russian seas) reported a 
usual depth range of 25-100 m, Naumov giving extreme limits of 5-250 m. Kramp (1943, 
Greenland) gave a range of 25-650 m. 

Hincks (1868) stated the substrate to be other hydroids. All the BMNH material is on 
hydroids, especially Tubularia larynx (sens, auct., e.g. Hincks, 1868), Hydrallmaniafalcata 
(Linnaeus, 1758) and Abietinaria abietina (Linnaeus, 1758), and other sertulariids. 
Although Couch (1844) reported material on the antennae of crabs and on a bivalve (Pinna 
fragilis, as '.P. ingens'), his description of the gonotheca suggests his material was Clytia 
hemisphaerica. He evidently confused the two species. 


REMARKS. Some nomenclatural confusion between the present species and Sertularia uni- 
flora Pallas, 1766, is discussed below (pp. 77-78). 

Rees & Thursfield (1965) suggested that C. volubilis might prove conspecific with 
Rhizocaulus verticillatus (p. 67). Their evidence was some similarity in the hydrothecae 
and gonothecae of the two species. However, the long BMNH series confirms the several 
constant differences. The linear dimensions of hydrothecae, hydrothecal pedicels, gono- 
thecae and stolon diameters in C. volubilis are about half the same dimensions in R. 
verticillatus. And while none of the C. volubilis specimens has polysiphonic, erect stems 
these are present in all the R. verticillatus specimens. There is no intermediate material. 
Further, the perisarc tubes in R. verticillatus are nearly all parallel. Had the two forms been 
conspecific, colonies of C. volubilis s. str. with some aggregation of the perisarc tubes might 
have been found; but there are no such specimens in the BMNH series. 

As noted by Hincks (1868) the distinction between C. volubilis and the hydroid stage of 
Clytia hemisphaerica was overlooked by Johnston (1847), but recognized soon afterwards by 
Alder (1 857). Couch ( 1 844) also confused the two. 

Confusion between C. volubilis and the species now called Calycella syringa (Linnaeus, 
1767) occurred in the mid-eighteenth century but was resolved by Linnaeus (1767) himself. 
Essential details are given in the above synonymy, and further discussion in Cornelius 
(1978). The species is currently referred to the family Campanulinidae. 

Campanularia groenlandica Levinsen, 1893, although widely recognized, was apparently 
founded on C. volubilis material from the Davis Strait. Syntype material in the BMNH shows 
features present in the original illustrations of groenlandica, for example spirally sculptured 
hydrothecal pedicels and blunt cusps on the hydrothecal rim; and the gonotheca shown in 
the original illustration is identical with that normal in C. volubilis. Thus the two taxa 
appear conspecific. Material has been recorded as C. groenlandica from Trondheim Fjord 
(Mathiesen, 1928) north to 68 20' N (Kramp, 1943; 50-525 m depth); and Naumov (1969) 
recorded 'C. groenlandica' from the N coast of Russia. ' 

The type material of Clytia mollis Stechow, 19190, examined here, comprises immature 
colonies of C. volubilis. Stechow's original illustration incorrectly shows a truncate, wide 
mouthed gonotheca. It is simply a young one in which the long neck has yet to form. The 
hydrothecal pedicels illustrated are topped by sub-hydrothecal spherules not present in 
Clytia. Stechow tentatively included in the synonymy of 'C mollis' the nominal species 
Clytia laevis Weismann, 1883, based on Naples material. However it is clear from 
Weismann's description that C. laevis was founded on normal Clytia hemisphaerica 
material, and it is here referred to that species. 

Genus ORTHOPYXIS Agassiz, 1862 

Clytia: Westendorp, 1843 : 23 (part; see Remarks under Orthopyxis Integra). 

ISilicularia Meyen, 1834 : 206 (?part; see Remarks and p. 50). 

Campanularia: Macgillivray, 1842:465 (part); Couch, 1844:40 (part); Hincks, 1868: 160 (part); 

Millard, 1975 : 203 (part); (see Remarks below, and under O. Integra). 
Clytia (Orthopyxis) Agassiz, 1862 : 297. 
Clythia Agassiz, 1862 : pi. 28 (lapsus for Clytia). 

Orthopyxis Agassiz, 1862 : 355; Ralph, 1957 : 834; Arai & Brinckmann-Voss, 1980 : 101. 
Hincksia Agassiz, 1862 : 355 (sic). 
Eucopella von Lendenfeld, 1883a : 188. 

Agastra Hartlaub, 1897 : 452 (nom. mid.), 504; Kramp, 1961 : 160. 
Leptomedusa Browne, 1900 : 7 14 (see notes on Nomenclature under O. Integra). 

TYPE SPECIES. Clytia (Orthopyxis) poterium Agassiz, 1862; by monotypy; may be conspecific 
with O. integra. Orthopyxis was introduced by Agassiz as a subgenus of Clytia on page 297 of 
his work, comprising the 'new' species poterium alone; but on page 355 he upgraded it to 
genus. On that page he implicitly used the combinations 'Orthopyxis (Orthopyxis) 
poterium\ 'Orthopyxis (Campanularia) volubiliformis' and 'Orthopyxis (Laomedea) Integra' 


(of various authors). Thus poterium should rightly be taken as type species by monotypy 
of the subgenus Orthopyxis Agassiz, 1862. Nutting's (1915 : 63) designation of Campanu- 
laria caliculata Hincks, 1853, as type species must be disregarded since caliculata was not 
originally included. It was unfortunately repeated by Arai & Brinckmann-Voss (1980). 

DIAGNOSIS. Campanulariidae forming stoloniferous or short unbranched upright colonies; 
stolon anastomosing; true diaphragm absent; hydrotheca fundamentally radially symmetri- 
cal but often asymmetrically thickened; medusa reduced, lacking manubrium and tentacles, 
not feeding, believed facultatively retained in at least one species. 

REMARKS. The genera Silicularia Meyen, 1834, and Hypanthea Allman, 18760, are 
discussed on page 50. 

Ralph (1957) listed some works in which Orthopyxis was discussed in relation to 
Eucopella von Lendenfeld, 1883a (based partly on Campanularia bilabiata Coughtrey, 
1875); and other discussion was provided for example by Bale (1914), Nutting (1915) and 
Fraser (1918). The species Eucopella campanularia was described in greater detail in 
another paper (von Lendenfeld, 1883/?). Bale, Ralph and others referred Eucopella to 
Orthopyxis; but Hirohito (1969) held the two genera distinct on the presence or absence 
respectively of marginal vesicles in the (retained) 'eumedusoid'. He referred caliculata 
Hincks, 1853, to Eucopella, stating an intention to discuss the generic question later. 
However, caliculata is here referred to O. Integra; and generic separation seems unjustified. I 
provisionally follow several previous authors in regarding Eucopella congeneric. However, 
Kramp (1961) accepted the genus and included in it the two species bilabiata Coughtrey, 
1875, and crenata Hartlaub, 1901. He designated bilabiata type species. The two species 
Kramp included may not be distinct: they are treated here under O. crenata (p. 60). 
Kramp's generic separation, from the 'medusa' genus Agastra, was based on minor differ- 
ences in the medusa and is not upheld here. 

The genus Hincksia Agassiz, 1862, was proposed to accommodate solely the well 
described species Campanularia tincta Hincks, 186 la; but Bedot (1910:311), Stechow 
(1923a : 94) and Rees & Thursfield (1965 : 93) referred the genus to Campanularia auct. In 
proposing the genus Agassiz stated merely 'The genus Hincksia is characterized by its one- 
sided, ringled, fertile hydra', hardly an acceptable diagnosis. Nevertheless, type material of 
C. tincta is available (BMNH reg. no. 1899.5.1.219-220), as noted by Rees & Thursfield. 
Although dry the material shows the characters of the genus Orthopyxis Agassiz, 1862, and 
Hincksia and Orthopyxis (not Campanularia) can be regarded congeneric. Under the first 
reviser principle I retain Orthopyxis, which has been widely used, and suppress Hincksia, 
which has not. O. tincta is an Australian species and so is outside the present scope, but it 
may be noted in passing that the type material has an anastomosing stolon like Orthopyxis s. 
str. and a highly distinctive, closely ringed gonotheca. Useful synonymies of the species were 
provided by Nutting (1915), Stechow (19230) and Rees & Thursfield (1965), and a redescrip- 
tion by Stepanyants (1979). Campanularia tincta sensu Warren, 1908, is mentioned here 
under C. africana Stechow, \923d, a junior synonym of O. crenata (p. 60). 

The genus Agastra Hartlaub, 1897, was based on Helgoland material of the medusa stage 
of Orthopyxis integra (see also p. 67). Kramp (1961) resurrected the genus but there seems 
no doubt that its synonymy with Orthopyxis is justified. 

Orthopyxis crenata (Hartlaub, 1 90 1 ) 
(Fig. 5) 

^.Campanularia bilabiata Coughtrey, 1875 : 291-292, pi. 20, figs 46-49. 

^.Campanularia everta Clarke, 1876:251, 253-254, pi. 39, fig. 4; Garcia Corrales el at., 

1978 : 24-25, fig. 9 (syn. C. lennoxensis Jaderholm). 

Eucopella crenata Hartlaub, 1901 : 364-366, pi. 22, figs 27-31, 33-35; Hirohito, 1969 : 7, fig. 7. 
^.Campanularia lennoxensis Jaderholm, 1904/7 : 268-269, pi. 12, figs 4-5. 
Campanularia ?inlermedia Stechow, 19 \9a : 66-68, fig. V. 
^Orthopyxis delicata Trebilcock, 1928 : 3, pi. 2, fig. 1 ; Garcia Corrales el ai, 1978 : 22-23, fig. 8. 


Campanularia crenata forma intermedia: Picard, 195 la : 345. 

Campanularia crenata: Picard, 1955: 186; Millard & Bouillon, 1973:47^18, fig. 6B-F; Millard, 

1975 : 204-206, fig. 68A-F (?syn. Orthopyxis delicata Trebilcock, 1928); Garcia Corrales et al., 

1978:1 9-22, fig. 7; (non Allman, 1 8766). 
Orthopyxis crenata: Trebilcock, 1928:3; Ralph, 1957:838-840, fig. 6g-v (syn. O. formosa 

Trebilcock, 1928); Rees & Thursfield, 1965 : 104. 

TYPE MATERIAL AND LOCALITY. The species was based partly on material from French Pass, 
Bare Island, New Zealand, and partly on the original description of Campanularia bilabiata 
Coughtrey, 1875. I have located none of the type material. Ralph (1957) restricted the type 
locality to French Pass. 

MATERIAL EXAMINED. I have seen no Atlantic material of this species. 

DESCRIPTION AND IDENTIFICATION OF HYDROID STAGE. The lack of available material of this 
species and the taxonomic confusion surrounding the whole genus together make redescrip- 
tion difficult. The following identification notes are adapted from Ralph (1957), Millard & 
Bouillon (1973) and Millard (1975). Differing from O. Integra as follows: hydrothecal rims 
smooth through gently wavy to crenate, with 8-12 short rounded cusps, commonly varying 
within a colony (always smooth in O. Integra); hydranth with c. 14 tentacles (> 20 in O. 
Integra: Ralph, 1957; but see p. 40). Other reported differences seem invalid (but see 
Dispersive stage). 

Fig. 5 Orthopyxis crenata. Hydrotheca, Port Phillip, Australia, intertidal; 1959. 10. 1.1. 

Scale lO^m. 

Variation. Ralph (1957) and Millard (1975) indicated that the variation in O. crenata 
parallels that in O. Integra (p. 63). 

DISPERSIVE STAGE. A medusa. By homology with O. Integra it might be expected that the 
medusa is short lived and does not feed. Hirohito (1969) described newly released medusae. 
The umbrella was sub-spherical (0'5 mm high, 0'6 mm wide). There was a distinct velum, 4 
broad radial canals and 8 statocysts; but no tentacles or stomach. Published descriptions 
suggest that the medusa of O. Integra differs in being proportionately taller. 

REPRODUCTIVE SEASON. Fertile material recorded early March near Marseille (Stechow, 

DISTRIBUTION. From N coast of Spain (Garcia Corrales et al., 1978, as Campanularia everta) 
and Mediterranean Sea southwards (S France, Stechow, 1919a as C. intermedia; Picard, 
195 la, 19586; Millard, 1975; Algeria, Picard, 1955; S Spain, Garcia Corrales et al.}. Widely 


distributed in warmer parts of all oceans (Millard). Cape Verde Islands (Rees & Thursfield, 

HABITAT. On Posidonia (eel grass) and Bryozoa (Millard & Bouillon, 1973, Seychelles); 
intertidal to about 3 m (Millard, 1975, southern Africa); 1-20 m, Spain (Garcia Corrales et 
al., 1978). Campanularia africana sens. Buchanan (1957), possibly conspecific, came from 
14 m ofTGhana (see Remarks). 

REMARKS. Authors who have placed this species in the genus Campanularia have apparently 
overlooked the senior homonym Campanularia crenata Allman, 18766 (see p. 52). If the 
present species is again referred to Campanularia another specific name would be required, 
and one of the names discussed by Hartlaub ( 1 90 1 ) might be available. 

Hartlaub thought O. crenata (Hartlaub) close to Eucopella Campanularia von Lendenfeld, 
1883a, 6, and 'identical with' Campanularia bilabiata Coughtrey, 1875. Ralph (1957), 
however, treated 'Orthopyxis crenata' and 'Silicularia bilabiata' under different genera. In 
this Ralph was unwise since crenata Hartlaub was, in part, a nom. nov. for bilabiata. The 
name bilabiata might prove to be available for the present species but to avoid further 
confusion crenata is retained pending a review of the whole genus. See also the notes on 
Eucopella (p. 58). 

Picard (19586) recorded the nominal species Orthopyxis everta (Clarke, 1876, as 
Campanularia, based on Californian material) from 'the Mediterranean'. The original 
description resembles the present species, and the two might prove conspecific; but I have 
seen type material of neither. Ralph (1957) separated them on the structure of the gonotheca. 
If a synonymy were propsed everta might take priority for the present species but C. bilabiata 
Coughtrey is still older. Vervoort (1972 : 87) redescribed 'O. everta' recently and gave further 

The nominal species Campanularia ?intermedia Stechow, 1919a, was based on material 
from Marseille. I have not located type material but the vegetative characters given in the 
description seem identical with those of the present species as currently understood. As 
suggested by Stechow, and also by Garcia Corrales et al. (1978), C. lennoxensis Jaderholm, 
19046, is probably conspecific. 

Campanularia africana Stechow (1923d: 104, nom. nov. pro C. tincta sensu Warren, 
1908, from Natal; non C. tincta Hincks, 186 la, from 'Australia', see p. 58) was recorded 
from Takoradi, Ghana at 14m depth by Buchanan (1957). O. africana has been 
distinguished from O. crenata by Millard (1975), who redescribed both, mainly on 
gonothecal characters; and from the several reportedly endemic South African species she 
recognized on variations in these characters alone. However, the relatively poor original 
descriptions of most of the nominal species and the general taxonomic confusion in the 
genus make it unwise to accept Buchanan's record without further evidence. It is the only 
record of 0. africana from north of the equator. 

Discussion of the non- Atlantic nominal species of Orthopyxis having crenate hydrothecal 
margins was provided by Ralph (1957). 

Millard (1975) provisionally referred Orthopyxis delicata Trebilcock, 1928, to the present 
species; and it seems likely that O. delicata sensu Garcia Corrales et al. (1978; N & S Spain) is 

Orthopyxis Integra (Macgillivray, 1842) 
(Fig. 6) 

IClytia undulata Lamouroux, in Freycinet, 1 824 : 6 1 7-6 1 8, pi. 94, figs 4-5. 

Campanularia Integra Macgillivray, 1842 : 465; Johnston, 1847 : 109, pi. 28, fig. 2 (syn. C. laevis: 

Saunders, in Johnston, 1847); Hincks, 1868 : 163-164, pi. 31, fig. 1; Levinsen, 1893 : 168-169, pi. 

5, figs 14-18 (syn. C. caliculata Hincks; C. gracilis Allman, 18766); Broch, 1918 : 159-162 (syn. C. 

compressa Clarke; C. ritteri Nutting, 1901a); Vervoort, 1946^:274-276, figs 120-121 (syn. C. 

laevis Couch; C. caliculata Hincks; C. breviscyphia Sars; Clytia (Orthopyxis) poterium Agassiz; 

Laomedea repens Allman); Millard, 1975:208-211, fig. 69 (syn. C. caliculata Hincks; C. 


compressa Clarke; Agastra mira Hartlaub; Agastra rubra Behner; ?Campanularia gracilis: 

Stechow, 1925). 

Clytia ryckholtii Westendorp, 1 843 : 23-24, pi. 1 , figs e, f. 
Campanularia laevis Couch, 1844:42; Gosse, 1855:25; (non C. laevis Hartlaub, 1905=junior 


Capsularia Integra: Gray, 1848 : 86 (?syn. Campanularia laevis Couch). 
Capsularia laevis: Gray, 1848 : 87. 
Campanularia caliculata Hincks, 1853 : 178-179, pi. 5, fig. B; Hincks, 1868 : 164-167, pi. 31, fig. 

2 (syn. C. breviscyphia Sars; Clytia (Orthopyxis)poterium Agassiz. 
Campanularia breviscyphia Sars, 1857 : 158-159, pi. 1, figs 12-13. 
Clytia (Orthopyxis) poterium Agassiz, 1862 : 297-304. 
Clythia poterium Agassiz, 1862 : pi. 28, figs 1-20, pi. 29, figs 1-5. 
Orthopyxis poterium Agassiz, 1862 : 355. 

Clytia posterior Wright, 1862 : 308 (lapsus pro poterium Agassiz). 
Laomedea repens Allman, 1871: 49, fig. 20. 
?Eucopella Campanularia von Lendenfeld, 1883a : 186-189. 
Campanularia compressa Clarke, 1877:214, pi. 8, figs 5-6; Patriti, 1970:34-35, fig. 43 (syn. C. 

platycarpa Bale). 

Campanularia borealis Marktanner-Turneretscher, 1890 : 206. 
Campanularia integriformis Marktanner-Turneretscher, 1890 : 207, pi. 3, fig. 2. 
'A leptomedusan' Browne, 1897 : 832, pi. 49, figs 3, 3a. 
Agastra mira Hartlaub, 1897:452, 504-506, pi. 22, figs 5, 8-10; Mayer, 1910:234 (syn. 

Campanularia caliculata Hincks); Russell, 1953:303-306, pi. 19, fig. 1, text-figs 186-188 (syn. 

' Leptomedusa sp.' Browne; Campanularia caliculata Hincks). 

Agastra caliculata: Browne, 1900 : 714-715 (syn. A. mira Hartlaub; Leptomedusa Browne). 
Leptomedusa gen.? sp.? Browne, 1900 : 714. 
Campanularia calyculata: Goette, 1907:193-204, pi. 15, figs 313-325 (syn. Clytia poterium 

Agassiz). , 

?Agastra rubra Behner, 1914 : 393-398, pi. 7, fig. 6, text-figs 8-10. 

Orthopyxis compressa: Stechow, 1919a : 69, fig. Wa-b; Picard, 19516 : 1 10; Picard, 1958a : 2. 
Orthopyxis asymmetrica Stechow, 1 9 1 9a : 71-72, fig. Xa-e. 
Clytia rijckholtii Leloup, 1947 : 22 (unjust, emend, pro C. ryckholtii Westendorp). 
Orthopyxis caliculata: Ralph, 1957:838, text-figs 6a-f (syn. O. macrogona von Lendenfeld); 

Picard, 1958& : 191 (syn. Campanularia integriformis auct.; see Remarks). 
Orthopyxis integral Rees & Thursfield, 1965 : 103-104. 
Eucopella caliculata: Hirohito, 1969 : 6-7, fig. 6. 

NOMENCLATURE. An unjustified emendation of the nominal species name caliculata, to 
calyculata, was followed by several authors (listed in Bedot, 1918, 1925). 

Browne (1 897) described the medusa of the present species but did not identify it, calling it 
simply 'A Leptomedusa gen.? sp.?'. Evidently Browne did not then regard Leptomedusa a 
generic name; but he later (Browne, 1900) used it thus: 'Leptomedusa gen.? sp.?', in a formal 
synonymy under 'Agastra caliculata (Hincks, 1853)'. Nevertheless it seems in keeping with 
Browne's intentions not to regard Leptomedusa as part of nomenclature. 

The widely used species name Integra may prove to be threatened by an older but obscure 
name, undulata. 

TYPE LOCALITY AND MATERIAL. Mouth of River Don, Aberdeen, Scotland; on Tubularia 
indivisa Linnaeus, 1758; material not located. 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia laevis Couch, r844, neotype, 
proposed herein. 

Campanularia caliculata Hincks, 1853, colony on Laminaria sp. and the red alga 
Phycodrys rubens (L.) Batt. (det. J. H. Price), in spirit, nr Old Head of Kinsale, Co Cork, Eire, 
coll. R. Allman, syntype; 1853.4.7.16. Remainder of type series, from Pegwell Bay, E. Kent, 
England, coll. R. S. Boswell, not located. The type locality of C. caliculata was restricted to 
Pegwell Bay by Ralph (1957) but the Co Cork material remains part of the syntype series. 

Orthopyxis asymmetrica Stechow, 1919a, infertile material on 2 microslides, Marseille; 
Munich Zoological Museum. 



OTHER MATERIAL EXAMINED. BMNH collection, c. 50 specimens, mostly from the British 

DESCRIPTION OF HYDROID STAGE. Colony a creeping hydrorhiza with single, irregularly 
spaced hydranths and hydrothecae on long pedicels and, separately, subsessile gonophores 
and gonothecae. Hydrorhiza smooth but sinuous, walls almost unthickened but often 
(Ralph, 1957; Millard, 1975) with a flat lateral flange of perisarc; branched frequently and 
(e.g. BMNH 1922.3.6.170, 1962.11.7.9) occasionally anastomosing. Hydrothecal pedicels 
usually narrower than hydrorhiza; walls usually much thickened; typically grooved with a 
smooth spiral; often 2-3 widely spaced shallow annuli near top; sub-hydrothecal spherule 
present; hydrotheca cup-shaped, length : breadth ratio variable; base wide to narrow, walls 
almost straight and diverging in narrower-based specimens; often flared near rim; walls thin 
to very thick, but rim region almost always unthickened; amount of thickening varying 
between adjacent hydrothecae and within a single hydrotheca (Fig. 6); rim even; small 

Fig. 6 Orthopyxis Integra, (a-e) 15-25 m, Espegrend, W Norway, 13 April 1962; 1962.10.7.1 1. 
(a-b) adjacent hydrothecae with differing pedicel lengths, (c) gonotheca, sex unknown, (d-e) 
sub-hydrothecal spherule, (f-h) Knysna, Cape Province, Republic of South Africa, 
1922.3.6.170. Scales: (a-c, 500 //m; (d-e) 10 //m; (g) 10 //m; (h) 10 //m. 

spherical chamber formed basally within hydrotheca by internal ring of perisarc. Hydranth 
with 20-30 tentacles (histological details in Agassiz, 1862; Stefani, 1956, 1959; Kawaguti, 
1966; anatomical details of a possibly conspecific form in von Lendenfeld, 18836). 
Gonotheca ^ = 9; broad, roughly parallel sided; truncated and slightly narrowing above, 
tapering more or less abruptly below; length usually l^-2x breadth, occasionally 5-6x (e.g. 
Allman, 1871 : fig. 20, as Laomedea repens; Vervoort, 1946a : fig. 120); usually laterally 
flattened but sometimes circular in transverse section; walls of gonotheca often thickened, 
sometimes much so; smooth through sinuous to deeply grooved spirally; aperture distal, 


nearly as wide as maximum diameter of gonotheca. Pedicel short to absent, usually unringed 
and grading into base of gonotheca (but see Fig. 6). Blastostyle with one well developed 
medusa, whether retained or released, and a second basal bud the fate of which seems 
unrecorded (see Dispersive stage). Present evidence suggets that medusae of either sex are 
sometimes retained. Nematocysts described by Ostman (1979). 

Variation in hydroid stage. See also the comments of Ralph (1957 : 838) and Millard 
(1975:209). Even among the Campanulariidae O. Integra is unusually variable in 
morphology, and is unusual also in its habit of sometimes releasing and at other times 
retaining the medusa. Some of the variation may be genotypic, but the controlling factors are 
hardly known. Naumov (1969) referred colonies with thick walled hydrothecae to a variety, 
caliculata Hincks, 1853, which he considered grew only in strong currents; but although this 
relation seems logical he offered no proof. In his introductory sections (p. 123) he reported 
that hydrothecae of this species grow larger in cool water than in warm. 

DISPERSIVE STAGE. Basically a short lived medusa. But this is often retained (as a 
gonomedusa), when the planula is the only motile stage. The free medusa was perhaps first 
described by Hartlaub (1897) from Helgoland, and shortly after by Browne (1897) working 
independently in SW Ireland. But von Lendenfeld (1883a, b) had earlier described a closely 
similar nominal species which may prove identical, from Australia ('Eucopella campanu- 
laria"; see Remarks); and Agassiz (1862, as Clytia poterium) had still earlier described 
planula release from retained medusae. 

The medusa, when released, is degenerate and ephemeral. It lacks organs of feeding and 
survives only a few days. Umbrella height c. 1 mm, width c. 0*65 mm; jelly thick, velum 
broad; stomach, manubrium and mouth absent; four narrow radial canals each with lobed 
gonad midway along; no tentacles or marginal cirri (after Russell, 1953). Apparently only 
one medusa at a time is produced from each blastostyle. The medusae swim actively 
(Hartlaub, 1897) but are probably short lived since they are presumed not to feed. They are 
sexually mature on release and do not develop further (Russell, 1953). 

Giard (1898) has often been thought the first to have linked the medusa to its hydroid but 
von Lendenfeld's (18836) earlier work might have been on this species (see Remarks). Giard 
was certainly the first to record that the medusa is not always released. He has been 
misquoted but his paper was quite explicit. Giard thought that time of year influenced 
medusa release, and so did Behner (1914) who worked on the probably conspecific 
Mediterranean medusa Agastra rubra Behner, 1914. But Stefani (1959) recorded liberation 
in turbulent water and retention under calmer conditions. Millard (1975; pers. comm.), 
however, stated that medusa release had not yet been recorded in southern African popu- 
lations (see also Remarks). The factors influencing release are still unclear. 

The female gonophore was recorded by several of the earlier workers but the male was not 
described until the work of Stefani (1956) and Hamond (1963), again excepting the much 
earlier and largely overlooked work of von Lendenfeld (18836) on the possibly identical 
Australian populations. 

Some authors (Russell, 1953, quoted in Rees & Thursfield, 1965; Hamond, 1963) have 
interpreted the retained medusae as sporosacs, but current knowledge of the life cycle 
confirms that they are medusoid. Following Miller (1973) they can be called gonomedusae. 
The often reported 'second medusa bud' near the base of the blastostyle [e.g. von Lendenfeld, 
18836; Giard, 1898; Hamond, 1963 (c?); Hirohito, 1969 (9); Millard, 1975 (9); BMNH 
1915.3.6.12 (d 1 ); also in congeneric species, Ralph, 1957] corroborates Miller's theory of 
descent from a gonophore producing medusae. Evidently the ancestral form produced 
several medusae on each blastostyle but today only one is produced at a time. 

REPRODUCTIVE SEASON. Free medusae recorded May-November in British waters (Russell, 
1953); June-September in NW France (Teissier, 1965); December-February & July at 
Naples (Lo Bianco, 1909). Some authors, from Giard (1898) onwards, have considered that 
medusa release occurs only towards the end of the reproductive season and that gamete 
release from sessile medusae occurs earlier in the year; but precise dates are unrecorded. 


Teissier (1965) found reproductive structures on the hydroid stage from May to October in 
NW France. 

DISTRIBUTION. Nearly cosmopolitan, occurring in all oceans from the intertidal to a little 
below Continental Shelf depths (at least in cold seas); and from the tropics to latitudes as high 
as 76 40' N (Greenland). The species is one of the most widely distributed of all hydroids. 
Noteworthy records from the eastern North Atlantic include: E & W Greenland (Kramp, 
1929, 1943), N Norway (Mathiesen, 1928), Greece (Yamada, 1965), Black Sea (Manea, 
1972; possibly also Naumov, 1960, 1969, as Campanularia integriformis, see Remarks), 
Morocco (Patriti, 1970, as Campanularia compressa), Ghana (Buchanan, 1957), Senegal 
(Leloup, 1939), Cape Verde Is (Ritchie, 1907); and the range of the species extends at least to 
the southernmost tip of Africa (Millard, 1975). 

However, there are gaps in this wide distribution. Broch (1928) thought the species to be 
absent from the Kattegat, Skagerrak and Baltic, and Stechow (1927) from the Baltic alone; 
but Kramp (1935) and Jagerskiold (1971) recorded it from W Sweden. Apparently there are 
still no records from the Baltic Sea. The species evidently did not occur in the Zuider Zee 
(Hummelinck, 1936) and has probably never been reliably recorded from Dutch waters 
(Vervoort, 19460). However, it is sometimes washed ashore on the Belgian coast (Leloup, 

Similarly, Irish Sea and W Scottish records are few: Bardsey I, Wales (Knight-Jones & 
Jones, 1956), Isle of Man (Bruce, Colman & Jones, 1963), Isle of Cumbrae, Clyde Sea 
(Chopin, 1894; Rankin, 1901), 10m depth in Cregan Narrows, Loch Creran, Argyll (C. 
Edwards colln, pers. comm.). Chumley (1918) recorded no Clyde Sea material; and Stephens 
(1905) gave only a few Irish localities: Belfast, Dublin and Co Cork, the last including some 
of the syntypes of Campanularia caliculata. Possibly the only record from the west coast of 
Ireland is from Valencia I, Co Kerry (Browne, 1900), incidentally one of the earliest 
descriptions of the medusa. Hincks (1868) and Russell (1953) similarly listed no records from 
the western coasts of Eire and Scotland but the species is small and may have been over- 
looked. Broch (1918) included the NW Irish and W Scottish coasts in the North Atlantic 
distribution but did not cite material and may have been guessing. 

Lastly, Arai & Brinckmann-Voss (1980: 103) thought the species might not occur in 
British Columbia and Puget Sound. 

HABITAT. World depth data range from intertidal (e.g. Hincks, 1853, British Isles) through 
300 m (Broch, 1918, Davis Strait) exceptionally to 470 m (Kramp, 1929, W Greenland). The 
deepest records are from cold waters. The species has been recorded on a wide variety of 
algae, hydroids, other animals and inorganic substrates, and there is no regular association. 
Mathiesen (1928) recorded O. integra on Laminaria sp. to depths of c. 100 m off Norway. 
Broch (1918) regarded the species as stenohaline, a view supported by the lack of records 
from the Baltic Sea and Zuider Zee. 

REMARKS. Several species closely related to O. integra were recognized by Ralph (1957), 
Millard (1975) and Gow & Millard (1975), and a world revision of the genus would be 
timely. Much discussion was provided by Arai & Brinckmann-Voss (1980). 

O. integra itself is both variable and nearly cosmopolitan, and has consequently been 
described under many species names (cf. Clytia hemisphaerica, p. 73). The above 
synonymy includes only North Atlantic synonyms and main ones from other areas when 
they enter the discussion. The following notes on them are arranged in date order where 

Baster (1762 : pi. 2, fig, 7A, a) published some unidentified illustrations which Maitland 
(1876) referred to the present species. Vervoort ( 1 946a : 276) regarded them as indeterminate, 
however, and I concur. They are discussed further under Clytia hemisphaerica (p. 78). 
Although Pallas (1766) arguably applied an earlier species name partly to Baster's illustra- 
tions, the later name integra is not threatened (but see the paragraph after next). 

Clytia urnigera Lamouroux (1816: 203, pi. 5, fig. 6), based on 'Australasian' material, was 


discussed by Couch (1844) when proposing Campanularia laevis Couch, a nominal species 
considered below. Couch noted a resemblance between the hydrothecae of urnigera and 
those of laevis but rightly pointed out that the narrow gonothecal aperture of urnigera 
contrasted with the wide aperture in laevis, and held them distinct. C. urnigera is here 
referred to Clytia hemisphaerica (p. 78); and C. laevis to O. Integra (see below). 

Clytia undulata Lamouroux, in Freycinet, 1824, was based on a fertile colony growing on 
'marine plants' at Port Jackson, New South Wales. The species was regarded by Lamouroux 
as close to 'Clytia urnigera Lamouroux', here referred to C. hemisphaerica. It was mentioned 
again only twice in the literature according to Bedot (1905), in 1824 and 1836, but no further 
taxonomic features were mentioned. C. undulata seems to be closer to the present species 
than to 'C. urnigera' and C. hemisphaerica. It is mentioned here as it predates 
other Orthopyxis species and might prove conspecific with O. Integra; but before it can be 
fully assessed more information is needed about the Australian populations of Orthopyxis. 

Clytia ryckholtii Westendorp, 1843, was based on material from Ostend, Belgium. It was 
referred to O. Integra by Billard (1914); and also by Leloup (1947) as 'C. Rijckholtii Slab.\ 
Both Leloup's spelling and his reference to Slabber are wrong: Slabber (1769-1778) did not 
treat O. Integra or anything similar. I have not located the type material; but Westendorp's 
illustrations show a reptant colony with long hydrothecal pedicels spirally grooved top and 
bottom each with an intervening smooth portion, and an even rimmed hydrotheca. They are 
the earliest illustrations of O. integra yet identified but those of Clytia undulata 
Lamouroux, in Freycinet, 1824, which might prove conspecific, are earlier. The description 
of 'ryckholtii' states the pedicels to be 3 mm long and mentions an even rimmed hydrotheca, 
confirming the identification. A rare and later work by Westendorp (1853) on Belgian 
zoophytes was illustrated by actual specimens. Had C. ryckholtii been included the speci- 
mens might have been types; but it was not (see note 1, p. 123), and I agree with Billard 
(1914) that the type material is probably lost. 

Campanularia intertexta Couch, 1844, was based on a mixed type series comprising 
Lafoea dumosa (Fleming, 1 820) and an unidentified campanulariid which might have been 
O. integra. C. intertexta is now referred to L. dumosa (see p. 122) but the original 
illustration, which included the campanulariid, was one of the earliest of O. integra. 

Campanularia laevis Couch, 1844, type locality Polperro, Cornwall, was implicitly 
referred to the present species by Johnston (1847). The type material may be have been 
preserved for a long time in the Royal Institution of Cornwall, Truro, but if present would 
have been destroyed by a flood in 1953 (Curator, Roy. Inst. Cornwall, pers. comm.). 
Johnston (1847) mentioned material sent to him by W. W. Saunders (BMNH reg. no. 
1847.9.24.65, on a herbarium sheet). The material is labelled in Johnston's hand 
'Campanularia laevis! W. Wilson Saunders, Hastings, 1840' and a label has been glued on 
later, also in Johnston's hand, on which is written 'Campanularia integra'. I concur with 
Johnston's later identification as O. integra. In the absence of the original type material I 
designate the specimen 1847.9.24.65 neotype of C. laevis Couch, 1844; and extend the type 
locality to comprise coastal waters of the south of England. I agree also with Johnston's 
suggestion that the original description of C. laevis Couch should be referred to O. integra; 
and with the tentative but similar opinion of Bedot (1905 : 1 57) that the two are conspecific. 
Gray (1848 : 86) too referred both the species C. laevis and the material just mentioned to O. 
integra; but, inconsistently, on the next page gave C. laevis Couch full specific rank. He did 
this under the genus name Capsularia Cuvier, 1797, now regarded a junior objective 
synonym of Coryne Gaertner, in Pallas, 1774 (see Cornelius, \915b : 378). Turning to a later 
work, it seems that Hincks' (1868 : 164) Hastings record of Saunders' material refers to the 
same specimen. Since Johnston (1847) did not publish the locality it seem likely Hincks 
deduced if from Gray (1848), who did. The junior homonym Campanularia laevis, 
Hartlaub, 1905, is discussed under Campanularia hincksii, which that nominal species most 
resembles (p. 54). 

Campanularia caliculata Hincks, 1853, is the main European synonym to have been 
applied to the hydroid stage. When proposing the nominal taxon Hincks in fact provided the 


first good description of the present species. Bedot (1918) recorded that Levinsen (1893) was 
the first to refer caliculata to Integra. Several senior authors have accepted this synonymy 
(e.g. Broch, 1918; Kramp, 1935; Vervoort, 1946a; Millard, 1975) and examination of type 
and non-type BMNH material supports their view. Hincks (1853) stated clearly the 
differences from Integra as he saw them. They were simply: a 'double' hydrotheca and 
pedicel (inaccurate observation of thick walled material) and a more gradual tapering of the 
hydrotheca in caliculata than in Integra (a variable feature). Neither character is reliable (see 
the above description and the discussion by Millard, 1975 : 209-210). Russell (1953) drew 
attention to the gonothecal contents later illustrated by Hincks (1868 : pi. 31, figs, la, b) as 
Integra. They seemed to be sporosacs, and Russell was not entirely ready to accept the 
synonymy. It then seemed plausible that there were two species, one with sporosacs and the 
other with facultatively released medusae. The question was resolved when Millard (1975) 
illustrated structures similar to those shown by Hincks and described release of gametes from 
them. Millard had not recorded medusa release in her area (southern Africa). Still, she 
commented 'In partly spent gonophores the medusoid structure can sometimes be seen and 
is best observed by dissecting the gonophore out of the gonotheca' (op. cit., p. 209). Evidently 
the sessile eumedusoids (called gonomedusae by Miller, 1973) resemble sporosacs closely 
and their medusoid nature is not easy to see. Hence Russell's point is answered, and 
synonymy seems in order. 

Campanularia breviscyphia Sars, 1857, was referred to C. caliculata Hincks, 1853, by 
Hincks (1868), and to the present species by Vervoort ( 1 946a). 

Clytia (Orthopyxis) poterium Agassiz, 1862, was a nominal species apparently based on 
North American material. Agassiz' description was exceptionally detailed and beautifully 
illustrated, but he failed to consider the several related species already described from 
European waters (integra, ryckholtii, laevis, caliculata, breviscyphia). His account suggests 
strongly that his material was merely O. integra. Hincks (1868) and Bale (1914) summarily 
dismissed poterium as conspecific; but Agassiz' account remained the most detailed of the 
species for many years. It was the earliest of the medusa, albeit of retained specimens. The 
caption to the plates included the spelling ^Clythia\ an unjustified emendation of Clytia. 

Laomedea repens Allman, 1871, was referred to the present species by Bedot (1910) and 
Vervoort (1946a), and I concur. The originally illustrated material was said to have come 
from Scotland (Allman, op. cit., p. 48) and comprised only a female gonophore of O. 
integra. The nominal species was apparently never described again. Allman later applied the 
same specific name to another taxon, Hypanthea repens Allman, 1876a, type locality 
Kerguelen Island (see also p. 50). 

Campanularia gracilis Allman (18766 : 260, pi. 12, figs 5-6), from Japan, was referred to 
the present species by Levinsen (1893) but this was quite unjustified. The type material 
(BMHN reg. no. 1877.4.12.5) is clearly distinct and not closely related to O. integra. Stechow 
(1925 : 423, fig. 6) described new material of C. gracilis Allman but Millard (1975 : 208) 
provisionally referred it to O. integra. However, Stechow's illustration closely resembles the 
type material and Millard, like Levinsen, was apparently mistaken in uniting the two taxa. 
Yamada (1959 : 35) evidently regarded C. gracilis as valid but recorded no material apart 
from the type. 

Campanularia compressa Clarke, 1877, based on the hydroid stage, was referred to the 
present species by Broch (1910, 1918) and Millard (1975), although Arai & Brinckmann- 
Voss (1980) regarded it distinct. Other material of O. integra was recorded under the species 
name compressa by Stechow (1919a, Villefranche), Picard (19516, Senegal), Picard 
(1958#, Israel) and Patriti (1970, Morocco). See also the comments on Agastra rubra 
Behner, 1914, the corresponding nominal species based on the medusa, below. 

Eucopella Campanularia von Lendenfeld, 1 883#, is discussed above (p. 60). 

Campanularia borealis Marktanner-Turneretscher, 1890, was based on hydroid material 
from Spitzbergen. No illustration was provided. Linko (191 1 : 164) referred the species to O. 
integra and I concur. 

Campanularia integriformis Marktanner-Turneretscher, 1890, based on hydroid material 


from Trieste, seems the same as O. Integra. Naumov (1960, 1969) briefly described speci- 
mens from the Black and Adriatic Seas as C. integriformis but this too was probably O. 
Integra. If so, Naumov's report of 0. Integra in the Black Sea predates that of Manea (1972), 
who had claimed the first record. Picard (1958&) referred integriformis to caliculata Hincks, 
1853, without comment but caliculata, too, is now referred to Integra. 

Agastra mira Hartlaub, 1897, based on Helgoland material, has been widely regarded as 
the first description of the free medusa of O. Integra. Browne (1897) described the medusa 
from SW Eire in the same year, as an unidentified leptomedusan; but a footnote in Browne's 
paper comments on Hartlaub's account, which had presumably already appeared. Both 
accounts of the medusa might be predated, however, if the closely similar Australian 
nominal species Eucopella campanularia von Lendenfeld, 1883#, b, proves conspecific. 

Several authors listed by Bedot (1918, 1925) applied the combination Agastra caliculata to 
the medusa stage. 

Agastra rubra Behner, 1914, a medusa from the Mediterranean, was referred to O. integra 
by Stefani (1959) and Millard (1975) among others. Its nominal hydroid, Campanularia 
compressa Clarke, 1877 (see above), was referred to O. integra by Broch (1910, 1918); and 
both stages were so treated by Millard (1975). Kramp (1961), however, considering the 
medusa stage alone, held A. rubra distinct on the shape of the gonads. Further work seems 
necessary to resolve these different views, but the majority opinion is that A. rubra is invalid. 

Finally, the type material of Orthopyxis asymmetrica Stechow, \9\9a, from Marseille, 
was examimed and found merely to be O. integra. Philbert (1935a) described growth forms 
under this name. 

Genus RHIZOCAULUS Stechow, \9\9b 

Sertularia Linnaeus, 1758 : 81 1 (part). * 

Campanularia Lamarck, 1816 : 112 (part); Hincks, 1868 : 160 (part). 
Rhizocaulus Stechow, 19 196 : 852; Stechow, 1919c: 16. 
Verticillina Naumov, 1960 : 9, 1 15, 122, 269; Naumov, 1969 : 6, 1 15, 123, 291. 

TYPE SPECIES. Sertularia verticillata Linnaeus, 1 758; by original designation. 

DIAGNOSIS. Campanulariidae forming erect, polysiphonic colonies; hydrothecae in whorls; 
no true hydrothecal diaphragm; sub-hydrothecal spherule present; no medusa stage. 

REMARKS. Nutting (1915 : 28) designated Sertularia verticillata Linnaeus, 1758, type species 
of the genus Campanularia Lamarck, 1816, but for the reasons given under that genus 
(p. 5 1) I have applied to the International Commission on Zoological Nomenclature for that 
designation to be set aside (Cornelius, 1981). 

Rees & Thursfield (1965) opposed the erection of a distinct genus to accommodate 
verticillata on the grounds that in Lafoea Lamouroux, 1821, family Lafoeidae, there are 
both stolonal and erect colonies; but colony habit does not have the same taxonomic value in 
all families, and the separation is upheld here. 

Stechow (1919/7) listed several nominal species in the genus but they may prove 
conspecific with R. verticillatus. 

Rhizocaulus verticillatus (Linnaeus, 1758) 
(Fig. 7) 

Corallina ramosa, ramis singulis equisitiformibus in summis capillamentis contortis et verticillatim 

dispositis . . . Ellis, 1 755 : 23-24, pi. 1 3, figs A, a. 
Sertularia verticillata Linnaeus, 1758 : 81 1. 
Campanularia verticillata: Lamarck, 1816: 113; Hincks, 1868: 167-168, pi. 32, fig. 1, la; Goette, 

1907: 179-189, pi. 14, figs 294-304, pi. 15, figs 305-306; Linko, 1911 : 188-200, fig. 35 (syn. Clytia 

olivacea Lamouroux, 1 82 1 ). 

Clytia olivacea Lamouroux, 1821 : 13, pi. 67, figs 1-2. 
Campanularia verticellata Couch, 1842 : 49 (lapsus). 



Campanulata verticillata: Agassiz, 1862 : 354, footnote (unjustified emendation of Campanularia). 
Rhizocaulm verticillatus: Stechow, 19196 : 852: Stechow, 1919c : 16; Stechow, 1923a : 105-106. 
Verticillina verticillata: Naumov, 1960 : 269-270, fig. 159; Naumov, 1969 : fig. 159. 

TYPE MATERIAL AND LOCALITY. Linnaeus (1758) provided only a diagnosis of this species, 
implying that he did not see material (cf. Cornelius, 1979 : 309). Indeed, there is none in the 
Linnaeus collection held by the Linnean Society of London (Savage, 1945). Linnaeus (1758) 
cited only the illustration of Ellis (1755 : pi. 13, fig. A, but not fig. a), and the colony Ellis 
illustrated can be regarded as holotype. It was collected from the coast of Cumberland, NW 
England, to which the type locality can be restricted. The specimen is almost certainly now 
lost (Cornelius, 1975a : 267, footnote). 

MATERIAL EXAMINED. BMNH collections, about 40 specimens. Some northerly material 
deserves mention: Barents Sea, 74 8' 50" N, 30 31' 28" E, 375m, 1882, ex D'Arcy 
Thompson collection; 1956.1 0.23 . 1 80. 

DESCRIPTION. Colony large, erect, occasionally and irregularly branched; stems polysiphonic 
with pedicels roughly in whorls, recalling the terrestrial horse-tail plants (Equisetum L.). 
Component perisarc tubes straight, parallel, each bearing straight, smooth or spirally 
grooved hydrothecal pedicels (sometimes annulated, e.g. Fig. 7) at approximately regular 
intervals. Hydrotheca bell shaped, sub- hydrothecal spherule present; rim with c. 12 blunt 
cusps. Gonotheca ?rf = 9, flask shaped, with neck of varied length; on short pedicel. 

Fig. 7 Rhizocaulus verticillatus. (a) part of colony, Isle of Man, 25 March 1894; 1948.10.1.21. 
(b) gonotheca with long neck, 30-40 m depth, nr Bergen, 15 August 1962; 1962.11.7.8. (c) 
gonotheca with short neck, 45 m depth, W Scotland; 1 888.4.2.39. Scale (a-c) 500 //m. 

DISPERSIVE STAGE. Planulae, probably developing within the gonotheca. Early gonophore 
development was described by Goette (1907). No medusa. 

REPRODUCTIVE SEASON. Fertile material recorded May, 1934, off NE England (H. O. Bull, in 
Evans, 1978); July, NW France (Teissier, 1965); 15 August, 1962, nr Bergen, Norway 
( 1962.11.7.8). 


DISTRIBUTION. Widespread in sublittoral and coastal waters from Tromso, Norway 
(Mathiesen, 1928) and Danmarks Havn, Greenland (Kramp, 1943) at least to Roscoff, NW 
France (Teissier, 1965). The species is well known from offshore areas of Britain, the 
Netherlands, Belgium, Denmark and Sweden. See also Material examined. 

The status of JR. verticillatus along the W coast of France and the Bay of Biscay is unclear. 
Vervoort (\946a) and Naumov (1969) recorded it but the species was not listed in several 
local faunal surveys (Billard, 1927; Nobre, 1931; Da Cunha, 1944, 1950; Fey, 1969) and 
there is no BMNH material from south of the English Channel. Picard (19586) did not 
record the species from the Mediterranean Sea; but Manea (1972) provided an acceptable 
record from the Black Sea, possibly the farthest south the species has been found on 
European coasts. 

HABITAT. Naumov (1969) gave a usual depth range of 50-200 m, with extremes of 15 m and 
680 m. Mathiesen (1928) gave a range of 20 m to 600 m in S Norway. 

Hamond (1957) associated the species with 'sandy grounds' off the Norfolk coast, but 
Teissier (1965) recorded it on algae in NW France. Hincks (1868) and Leloup (1952) found 
the species on pebbles and shells. R. verticillatus seems to occur typically on occasional solid 
substrates in otherwise sandy areas. 

REMARKS. There seems no need for revision of this distinctive species. Rees & Thursfield 
(1965) drew attention to the superficial similarities between it and Campanularia volubilis 
(but see p. 57). 

The development and structure of the compound stem of R. verticillatus was studied by 

The combination Campanularia verticillata var. grandis Hickson & Gravely, 1907, 
related to an antarctic species now known as Stegella grandis (Hickson and Gravely) and 
assigned to the Campanulinidae (e.g. Totton, 1930; Stepanyants, 1979). The similarity in 
colony form is striking but S. grandis lacks a sub-hydrothecal spherule and the hydrothecal 
rim is quite different. 

Subfamily CLYTIINAE nom. nov. 

Phialidae Haeckel, 1879 : 163 (part). 
Phialinae: Mayer, 1910 : 232 (part). 
Campanularinae: Russell, 1953 : 284. 

DIAGNOSIS. Campanulariidae with a usually reptant, branched but not anastomosing stolon; 
with true hydrothecal diaphragm; usually without subhydrothecal spherule (but present in 
Clytia hummelincki); medusa present in some genera, a typical leptomedusa with prominent 

SCOPE. The genera Clytia Lamouroux, 1812; Gastroblasta Keller, 1883; and Tulpa Stechow, 
192 la. Gastroblasta and Tulpa have not been recorded in the eastern North Atlantic and are 
discussed only under Remarks, but Clytia is treated below. 

TYPE GENUS. Clytia Lamouroux, 1812. 

REMARKS. Previous subfamily names applied in part to this group (Phialidae Haeckel, 1879; 
Phialinae Mayer, 1910) have been derived from Phialium Haeckel, 1897, the name of a 
genus now referred to the Lovenellidae (Mayer, 1910; Kramp, 1961). Phialium can be taken 
as type genus of the nominal subfamily Phialiinae; and there seems no available name for the 
present group. 

There has no doubt been confusion between Phialium Haeckel and Phialidium Leuckart, 
1856. Phialidium is today referred to Clytia Lamouroux, 1812, and basing a new subfamily 
name on Phialidium might be confusing. Haeckel (1879 : 186) indicated that he understood 
the relation between Phialidium and Clytia by including Clytia johnstoni sensu Bohm in the 


synonymy of P. variable. This relationship had only just been understood (p. 76) and 
Haeckel might have been reticent to consider all the nomenclatural implications. 

The subfamily name Clytiinae is to be considered a nom. nov. applied to the present sub- 
family as restricted by Russell (1953) under the preoccupied name Campanulariinae. It 
should not be confused with family group names derived from Clytus, a coleopteran genus 
name. These are spelt Clytinae, Clytini and so on (C. R. Smith, pers. comm.), being derived 
from the root Clyt. The root from which Clytiinae is derived is Clyti. 

The genus Gastroblasta Keller, 1883 : 622, is now restricted to a single, distinctive medusa 
species based on Red Sea material (Kramp, 1961). The name was once applied to species 
now referred to one or other of the Clytia species treated below (discussion on p. 72). 

The genus Tulpa Stechow, 1 92 la : 254, was proposed to embrace three species none of 
which has been recorded from the eastern North Atlantic. The type species is Campanularia 
tulipifera Allman, 1888, by original designation (= C. tulpifera lapsus auct.). The genus was 
accepted by Totton (1930) who described another species. Tulpa was reviewed by Ralph 
(1957) and redefined by Stepanyants (1979); and discussed also by Vervoort (1972) and 
Gravier-Bonnet (1979) who gave systematic notes. 

Genus CL YTIA Lamouroux, 1812 

Medusa: Linnaeus, 1767 : 1096 (part). 

Sertularia Ellis & Solander, 1786:51 (part). 

Oceania Peron & Lesueur, 18100 : 343 (part). 

Clytia Lamouroux, 1812 : 184. 

Campanularia auct., part (see Remarks under Clytia hemisphaerica, p. 77). 

Thaumantias Eschscholtz, 1829 : 102; Forbes, 1848 : 52 (part). 

Silicularia Meyen, 1 834 : 206 (part; see p. 50). 

Eucope Gegenbaur, 1856 : 241 (part). 

Phialidium Leuckart, 1856 : 18; Kramp, 1961 : 164. 

Epenthesis McCrady, 1857 : 191; Haeckel, 1879 : 182. 

Clytia (Trochopyxis) Agassiz, 1862 : 304. 

Clytia (Platypyxis) Agassiz, 1862 : 306. 

Clytea Wright, 1862 : 308 (lapsus pro Clytia). 

Clythia van Beneden, 1 866 : 166 (lapsus pro Clytia). 

Gastroblasta auct., part (non Keller, 1 883, s. str.). 

Pseudoclytia Mayer, 1900 : 53. 

Multioralis Mayer, 1900 : 54. 

non Clythia Agassiz, 1862 : pi. 28 (lapsus pro Clytia, but referred to Orthopyxis, p. 57). 

TYPE SPECIES. There were three species originally included in the genus, listed thus: 
'Sertularia volubilis Ellis, S. syringa Ellis, S. verticillata Ellis'. The descriptions cited are 
presumably those of Ellis & Solander (1 786), in which book binominals were employed, and 
not those of Ellis (1755) which lacked them. The point is important since 'S. volubilis sens. 
Lamouroux, 1812' was subsequently designated type species (see next paragraph). Confusion 
was caused when Lamouroux himself (in Lamouroux, Bory de Saint-Vincent & 
Deslongchamps, 1 824 : 202) later applied the name S. volubilis to another species, citing 
therein the Ellis, 1 755, illustration which in fact shows the species here called Campanularia 
volubilis (p. 55). Lamouroux' (1812) earlier citation, then, refers to Sertularia volubilis Ellis 
& Solander, 1786 [ = the nominal hydroid species Clytia johnstoni (Alder, 1856#), usually 
regarded the same as the medusa Clytia hemisphaerica Linnaeus, 1767; see p. 77]; and the 
later citation (Lamouroux et al., 1824), employing the same combination, refers to 
Campanularia volubilis (Linnaeus, 1 758) (see p. 77 for further discussion). 

The type species of Clytia might appear to be Sertularia volubilis Ellis & Solander, 1 786 
(non Linnaeus, 1758, see above), as designated by Mayer, 1910 : 262; but as just explained 
the combination is a junior homonym. This would be unimportant to current nomenclature 
if the hydroid Sertularia volubilis Ellis & Solander, 1786, were definitely known to be the 
same species as Medusa hemisphaerica Linnaeus, 1767. A relation is usually assumed (see 


notes under Clytia hemisphaerica, p. 79), and Linnaeus indicated Gronovius' (1760 : pi. 4, 
fig. 7) illustrated specimen, collected from Belgian waters (the type locality of hemi- 
sphaerica}. The illustration shows a strongly convex exumbrella surface, suggesting C. 
hemisphaerica sens. auct. (e.g. p. 73); but the lingering doubts over the identity of the 
hydroid C. gracilis (Sars, 1850; see p. 78) make the relation of the two nominal species 
uncertain. Therefore, it is useful to establish a soundly based name for the type species of the 
genus Clytia. The earliest available name which can be unequivocally linked with S. 
volubilis sens. Ellis & Solander, 1786, is Campanularia johnstoni Alder, 18560. Hence the 
correct name for the type species of Clytia is C. johnstoni. The often quoted synonymy with 
C. hemisphaerica, repeated below, is subjective. See also Cornelius (1981), and page 78 
concerning the very similar 'Laomedea gracilis' Sars, 1850. 

DIAGNOSIS. Colonial Campanulariidae with free medusa stage; hydroid generation forming 
umbranched stoloniferous or branched upright colonies; hydrotheca with true diaphragm, 
rim sinuous to deeply indented with round to sharp clefts and cusps; diaphragm transverse; 
no sub-hydrothecal spherule; medusa hemisphaerical to flat, with hollow marginal tentacles, 
velum well developed; manubrium short; 4-8 marginal tentacles on release (in species so 
far studied), many tentacles in adult. 

REMARKS. A summary of the species of Clytia in which the life cycles have been worked out 
was given by West & Renshaw (1970); and Roosen-Runge (1970) and Arai & 
Brinckmann-Voss (1980) gave detailed appraisals of several North American species. 

Early generic names applied to the type species were broad in concept (Sertularia 
volubilis = Medusa hemisphaerica = Campanularia johnstoni) and the diagnosis of them by 
successive workers often overlapped. Hence it is not unexpected that species now included in 
Clytia s. str. should at one time or another have been included in one or more other genera. 
There is little value in discussing past uses of generic names such as Medusa, Sertularia, 
Oceania and Campanularia for species now referred to Clytia. 

The name Thaumantias Eschscholtz, 1829, was also once widely used. It is a junior 
subjective synonym of Clytia, since Forbes (1848 : 41) nominated Medusa hemisphaerica 
Linnaeus, 1767, its type species. This medusa is, of course, widely held to come from the 
hydroid which is type species of Clytia; but the link is neverthless subjective and is likely to 
remain so as the original description of the medusa was brief. 

The genus Silicularia Meyen, 1834, is discussed on page 50. 

Availability of the generic name Eucope Gegenbaur, 1856, was discussed by Haeckel 
(1879), Mayer (1910), Rees (1939) and Russell (1953) among others. The four originally 
included species were referred to Obelia and Clytia (or Phialidium) by Mayer and 
subsequent authors; but so far as I can determine no type species has been designated. Russell 
(1953) referred three of the originally included species (E. thaumantoides, E. campanulata & 
E. affinis) to Clytia hemisphaerica (as Phialidium); and the remaining one (E. polystyla) is 
Obelia (e.g. following Cornelius, 19750). I designate E. affinis Gegenbaur, 1856, type species 
of Eucope, which falls in the subjective synonymy of Clytia (see also Bedot, 1910 : 414 and 
Hincks, 1868: 143). 

The genus Phialidium Leuckart, 1856, was based on a single species, P. viridicans 
Leuckart (1856 : 18-19, pi. II, figs 12, 14) which is therefore type by monotypy. Mayer 
(1910) confirmed its type status, and Kramp's (1961) designation of 'P. hemisphaericum L.' 
is incorrect. Russell (1953) drew attention to the close similarity between P. viridicans 
and Clytia hemisphaerica, including them in the same species synonymy. Thus the case for 
regarding Clytia and Phialidium congeneric is strong and Phialidium, being the later name, 
need no longer be used. 

The genus Epenthesis McCrady, 1857, was referred to Clytia by Mayer (1910 : 261) and 
Nutting (19 15 : 1) among others. The name is clearly a junior synonym of Clytia. 

The subgenus Platypyxis Agassiz, 1962, was referred to Clytia by Bedot (1910), and I 
concur (see Remarks under C. hemisphaerica). 


The genus Gastroblasta Keller, 1883, now includes just one species, from the Red Sea. All 
other uses of the generic name have related to abnormal medusae of Clytia species (Kramp, 
1 961; also Mayer, 1910, Kramp, 1957,1965). 

The genera Multioralis Mayer, 1900, and Pseudoclytia Mayer, 1900, were referred by 
Kramp ( 1 957, 1 96 1 ) to Phialidium Leuckart, 1856, and hence fall into Clytia. 

Luminescence has long been known in the genus (e.g. Darwin, 1860 : ch. 2, hydroid stage; 
other references in Forbes, 1848, medusa stage). Light emission was probably first recorded 
in Clytia by Macartney (1810). In October, 1804, he demonstrated to an invited audience at 
Herne Bay, Kent, that flashing in the medusa of C. hemisphaerica (which he called Medusa 
lucidd) was induced by raised temperature, electric shock and alcohol. His published 
illustration of the medusa is among the earliest of the genus. 

Clytia discoida (Mayer, 1900) 
(Fig. 8) 

Oceania discoida Mayer, 1900 : 5 1 , pi. 20, figs 53-55. 

Phialidium discoidum Mayer, 1910:272, pi. 33, figs 9-11; Kramp, 1959: 148, fig. 187; Kramp, 
1961 : 1 65-1 66; Schmidt & Benovic, 1977:637. 

TYPE LOCALITY. Tortugas, Florida; in plankton. 

DESCRIPTION (after Mayer, 1910; Kramp, 1959; Schmidt & Benovic, 1977). Adult medusa 
'quite flat' (Mayer), about 4 mm diameter; 1 6 short marginal tentacles with large basal bulbs; 
usually 3 statocysts between tentacle bases; velum well developed; 4 straight radial canals; 
gonads along almost whole length, eggs in 9 unusually large and prominent; manubrium 
'urn-shaped' (Mayer) with 4 recurved lips. Hydroid stage not recorded. 

Fig. 8 Clytia discoida. Adult medusa. Diameter c. 4 mm. Redrawn after Mayer (1910 : pi. 33, 

fig. 10). 

DISPERSIVE STAGE. The species is known from the medusa alone. See also notes under 
Dispersive stage of C. linearis. 

REPRODUCTIVE SEASON. Summer; June-August (Mayer, 1900; Schmidt & Benovic, 1977). 

DISTRIBUTION. Recorded from: Florida, Mexico, Brazil and W. Indies (references in Kramp, 
1961); southern Adriatic Trough (once only, Schmidt & Benovic, 1977). Several records 
from the Pacific Ocean were discredited by Kramp ( 1 96 1 ). 

HABITAT. Coastal plankton. 

REMARKS. The species is distinguished by its small size at maturity and by the extension of 
the gonads along almost the entire length of the radial canals. Kramp (1959, 1961) seems 
mistaken in stating the umbrella to be 'almost hemispherical'. Others have reported it nearly 
flat in the adult (Mayer, 1900, 1910; Schmidt & Benovic, 1977). However, younger stages 
are reportedly less flat (Mayer), but Schmidt & Benovic reported a young specimen only 


1*5 mm in diameter which was already 'more flat than hemispherical'. The adult diameter is 
4 mm. 

The species is known in the NE Atlantic from a single Adriatic specimen (Schmidt & 
Benovic, 1977). Kramp (1961) provided a literature synopsis. 

Clytia hemisphaerica (Linnaeus, 1 767) 
(Fig. 9) 

Medusa hemisphaerica Linnaeus, 1767 : 1098; Miiller, 1776 : 233. 

Sertularia uniflora:E\\\&, 1768 : 434, pi. 19, fig. 9. 

Sertularia volubilis: Ellis & Solander, 1 786 : 5 1 , pi. 4, figs E, e, F, f (non S. volubilis Linnaeus, 1 758; see 

p. 76 and Remarks). 

Oceania flavidula Peron & Lesueur, 1810a : 345; Peron & Lesueur, 718106 : 33. 
Oceania hemisphaerica: Peron & Lesueur, 18100 : 347; Peron & Lesueur, 18106 : 35. 
Clytia urnigera Lamouroux, 1816 : 203, pi. 5, fig. 6 (see Remarks under Orthopyxis Integra). 
Thaumantias inconspicua Forbes, 1848 : 52, pi. 8, fig. 3 a-d; Ritchie, 1911:31. 
ILaomedea gracilis Sars, 1850 : 138; Sars, 1857 : 160, pi. 2, figs 1-3, 5 (but not fig. 4, = Gonothyraea 

loveni; see Stechow, 19230 : 111); (non Dana, 1846 = Obelia dichotomd). 
Campanularia volubilis: (sens. Ellis & Solander) Hincks, 1852 : 84-85, pi. 3, fig. 5 (see Dispersive stage 

and Remarks). 

Campanularia sp. Gegenbaur, 1854 : 154, 189, pi. 1, figs 8, 8a, 9. 
Campanularia johnstoni Alder, 18560 : 359-360, pi. 8, fig. 8 (nom. nov. pro Sertularia volubilis Ellis & 

Solander, 1786). 

IPhialidium viridicans Leuckart, 1856 : 18-19, pi. 1, figs 12, 14 (see pp. 71, 86). 
IClytia noliformis McCrady, 1857 : 194-195, pi. 11, fig 4; Fraser, 1944 : 144-145, pi. 26, fig 1 17 (syn. 

C. simplex Congdon). 

Campanularia gegenbauriSars, 1857 : 48-49. 

Campanularia volubiliformis Sars, 1857 : 156 (nom. nov. pro Campanularia sp. Gegenbaur, 1854). 
Campanularia raridentata Alder, in Hincks, 18616 : 292; Hincks, 1868 : 176-177, fig. 2. 
Clytia (Trochopyxis) bicophora Agassiz, 1862 : 304-305, pi. 29, figs 6-9. 
?Clytia (Platypyxis) cylindrica Agassiz, 1 862 : 306-307, 354, text-figs 4 1-44, pi. 27, figs 8-9. 
Clytea vicophora Wright, 1862 : 308 (unjustified emendation of Clytia bicophora). 
Campanularia ?gigantea Hincks, 1866:297; Hincks, 1868:174-175, pi. 35, fig. 1; Jaderholm, 

1909: 19, 33, 69, pi. 7, figs 1-3. 
Clytia johnstoni: Hincks, 1868 : 143-146, pi. 24, figs 1, la (syn. Sertularia volubilis Ellis & Solander; 

Eucope campanulata Gegenbaur; E. thaumantoides Gegenbaur; E. affinis Gegenbaur; 

Campanularia gegenbauri Sars; Clytia bicophora Agassiz; see Remarks); Bohm, 1878 : 167-171, 

pi. 2, figs 1-9 (syn. Platypyxis cylindrica Agassiz; Eucope picta Keferstein & Ehlers, 1861; E. exigua 

Keferstein & Ehlers, 1861; E. variabilis Claus; E. gemmifera Keferstein; Thaumantias dubia 

Kolliker; T. thompsoni Forbes; T. convexa Forbes); Billard, 1928: 456-457 (syn. C. raridentata 

Hincks); Russell, 1953: 293, fig. 179; Ralph, 1957: 823-824, figs Ih-u, 3a-f(syn. C. compressa 

Totton, 1930). 

Clytia volubilis: DM Plessis, 1871 : 167-170, pi. 2; Hargitt, 1909 : 373-374. 
Clytia laevis Weismann, 1 883 : 1 58-1 59 (see Remarks under Campanularia volubilis, p. 57). 
Clytia flavidula: Metschnikoff, 18860 : 241-243, 257, 260, pi. 22, figs 9-10, 15. 
Campanularia? serrulata Bale, 1888 : 757, pi. 12, fig. 4. 

Campanularia raridentata var. Marktanner-Turneretscher, 1890 : 205, pi. 3, figs 3a-b. 
Thaumantias hemisphaerica: Browne, 1896 : 480-482 (syn. Medusa cymbaloidea Slabber, 1775; M. 

campanella Shaw & Nodder, 1795; M. lucida Macartney, 1810; Thaumantias lucida: Lesson, 1843; 

Epenthesis cymbaloidea Haeckel, 1879). 
Campanularia inconspicua: Calkins, 1899 : 349. 

^Campanularia attenuata Calkins, 1899 : 350, pi. 2, figs 9, 9a-c, pi. 6, fig. 9d. 
^Campanularia edwardsi Nutting, 19016 : 346, fig. 28. 
^.Campanularia pelagica Van Breemen, 1905 : 205-209, fig. 18. 
Clytia simplex Congdon, 1907 : 471-472, figs 14-15. 
Clytia obeliformis Stechow, 1914 : 128-129, fig. 6. 
IClytiapelagica: Billard, 1917 : 539-542, fig. 1. 
?Clytia serrulata: Stechow, 19190 : 46-47, fig. M. 


Campanularia acuta Stechow, 1919a:54 (nom. nov. pro C. raridentata var. Marktanner- 


Campanularia ?attenuata Stechow, 1919a : 61-62, fig. S (non Calkins, 1899). 
Campanularia brachycaulis Stechow, 1919a : 62-63, fig. T. 
Orthopyxis volubiliformis: Stechow, 1 9 1 9a : 70. 
Campanularia villafrancensis Stechow, 1919a : 157 (nom. nov. pro C. attenuata Stechow, 1919a 

(sic) : 6 1 , preoccupied). 

Clytia uniflora: Stechow, 1923a : 1 1 1 (non Sertularia uniflora Pallas). 
Thaumantias raridentata: Stechow, 1923a: 107-1 08, fig. 17. 
Clytia villafrancensis: Stechow, 1923a : 109-1 10. 
Clytia compressa Totton, 1930 : 146-148, text-fig. 6. 

tPhialidiumbicophorum.-T'hiel 1935 : 172; Kramp, 1959 : 149; Kramp, 1961 : 164-165. 
ILaomedea pelagica: Vervoort, 19460 : 285-288; Vervoort, 1959:313-315, fig. 55b-c; Vervoort, 

1968 : 15-17, fig. 5; Vervoort, 1972 : 91-92, fig. 26c. 
Laomedea gigantea: Leloup, 1952 : 161, fig. 93. 
Phialidium hemisphaericum: Billard, 1928 : 457 (syn. Thaumantias inconspicua Forbes); Kramp, 

1919 : figs 16-17 (syn. P. temporarium Browne, 1896; Thaumantias buskiana Gosse, 1853); Russell, 

1953 : 285-294, pi. 16, fig. 1, pi. 17, fig. 6, text-figs 172-179 (syn. Thaumantias pileata 

Forbes, 1841; T. sarnica Forbes, 1841; T. thompsoni Forbes, 1841; T. punctata Forbes, 1841; T. 

lineata Forbes, 1848; T. hemisphaerica: Forbes, 1848; T. inconspicua Forbes, 1848, Hincks, 1868; 7". 

buskiana Gosse, 1853; Eucope affinis Gegenbaur, 1856; E. campanulata Gegenbaur, 1856; E. 

thaumantias Gegenbaur, 1856; Phialidium viridicans Leuckart, 1856; ?T. typica Green, 1857; Clytia 
johnstoni: Hincks, 1868; T. leucostyla Spagnolini, 1876; Campanulina acuminata Bohm, 1878; 

Epenthesis cymbaloidea Haeckel, 1879; P. variabile Claus, 1881; Clytia flavidula Metschnikoff, 

1886; C. viridicans Metschnikoff, 1886; P. buskianum Browne, 1896; P. temporarium Browne, 1896; 

T. forbesi Johansen & Levinsen, 1903; Clytia volubilis: Sverdrup, 1921; for these references see 

Russell, 1953); Kramp, 1955 : 256-257 (syn. P. variabile Haeckel, 1879; ^Oceania languida Agassiz, 

Clytia hemisphaerica: Rees & Thursfield, 1965 : 95-96; Millard, 1966 : 478-480, fig. 14a-f; Vervoort, 

1968 : 16-17;Calder, 1975 : 300-302, fig. 4a-b; Millard, 1975 : 217-218, fig. 72a-d. 
Clytia gigantea: Rees & Thursfield, 1965 : 96-97. 
Thaumantias raridentata: Teissier, 1965 : 17. 

ILaomedea (Clytia) pelagica Garcia Corrales et al., 1978 : 28-29, fig. 1 1 . 
^Campanularia gracilis: (sensu Sars, 1850) Stepanyants, 1979 : 32, pi. 5, fig. 3 (syn. Clytia serrulata: 

Vanhoffen, 1910). 

IClytia sarsi Present paper, p. 78 (nom. nov. pro Laomedea gracilis Sars, 1 850, preocc; see p. 78). 
For further synonmy see Bedot (1901-1925), Russell (1953) and Kramp (1961). Kramp cited just the 
following combinations from the World literature not included in Russell's British list: Gastroblasta 
raffaelei, Clythia johnstoni, ^Phialidium languidum (provisionally retained distinct by Kramp), 
Pseudoclytia pentata f. hexaradiata. See also Clytia incertae sedis (p. 9 1 ). 

NOMENCLATURE. Millard (1966) has shown that the apparent use of the species name 
hemisphaerica by Gronovius (1760) was not strictly binominal. Neither also was that by 
Houttuyn (1770 : 423), cited by Bedot (1901 : 486) as Medusa hemisphaerica, despite the 
implication of Bedot's entry. Bedot dated Houttuyn's work as 1761-1773; but the volume in 
which the hemisphaerica reference appeared was published in 1770, three years after 
Linnaeus' (1767) introduction of the genuine binominal Medusa hemisphaerica. Linnaeus 
cited Gronovius' illustration as indication. 

The combination Clytia hemisphaerica results from the subjective linking of hydroid and 
medusa, and was probably first used by Rees & Thursfield (1965). The next year, Millard 
(1966) too discussed the combination. 

Many authors have placed hemisphaerica in the 'medusa' genus Phialidium Leuckart, 
1856; but Clytia Lamouroux, 1812, is older and now the life cycle is known should be used 
instead. The reason why Phialidium need not be used for any species is given above 
(P. 71). 

Browne (1896) incorrectly ascribed the combination Medusa hemisphaerica to Miiller, 
whose work Browne dated as 1766. The correct date of Miiller's work was 1776, and 
Linnaeus ( 1 767) has clear priority. 



TYPE LOCALITY AND MATERIAL. The type material of C. hemisphaerica Linnaeus, 1767, was 
the medusa described and illustrated by Gronovius (1 760 : 38, pi. 4, fig. 7). I have not tried to 
find it. Linnaeus gave the type locality as 'Belgian seas', from whence Gronovius' material 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. I have examined the type series of the hydroids 
Campanularia johnstoni Alder, 18560, and C. raridentata Alder, in Hincks 1861&, and 
consider them to be C. hemisphaerica (see Cornelius & Garfath, 1980; see also Remarks). 
The holotype colonies of the hydroids C. ?gigantea Hincks, 1866 (BMNH reg. no. 
1899.5.1.106, Lamlash, Arran, W Scotland) and Clytia obeliformis Stechow, 1914 (Munich 
Zoological Museum, fertile colony on microslide, Bergen) are also Clytia hemisphaerica. 

OTHER MATERIAL EXAMINED. This species is well represented in the BMNH collections. The 
following material, from other Museums, deserves mention: 53 01' N, 4 22' E, numerous 
colonies in spirit, coll. Lightship 'Texel', 1961, cf. 'Laomedea pelagica' (sensu Van 
Breemen), det. W. Vervoort; Rijksmuseum van Natuurlijke Historic, Leiden. 1040'N, 

Fig. 9 Clytia hemisphaerica. (a-b) 'pelagic' form, from microslide preparation in Institut Royal 
des Sciences Naturelles de Belgique, det. 'Laomedea gracilis" by A. Billard (mentioned, Billard, 
1917; 'probably ofTOstend'). (a) 'primary attachment disc' with four hydrothecal pedicels. Note 
absence of stolon, (b) hydrotheca. (c) 'pelagic form', gonotheca, 53 01' N, 4 22' E, det. 'Clytia 
pelagica" by W. Vervoort; colln Rijksmuseum van Natuurlijke Historic, Leiden, (d-e) 'benthic 
form'; (d) hydrotheca and (e) gonotheca, S England; 1934.8.17.19. (f) 'benthic form', base of 
pedicel branch from extremely sheltered locality, Caol Scotnish, Loch Sween, Argyll, W 
Scotland, 1 m, 30 May 1962; 1962.6.19.21. (g) 'benthic form 1 , base of pedicel branch, nr Bergen, 
40-90 m, 9 April 1962; 1962.10.7.19. (h-j) three hydrothecal rims from same colony, Scotland; 
1964.8.7.72. (k) hydrotheca, W Sweden, 20-30 m; 1962.11.8.10. (1) primary hydrotheca and 
attachment disc, reared from medusa ("Phialidium hemisphaericum* sens. Russell, 1953) by W. 
J. Rees, 2-17 March 1936; 1969.12.2.2. (m) medusa, Naples, redrawn after Mayer 
(1910: text-fig. 1 44). Scales: (a-c, d-1) 500 //m; (m) unknown. See also Fig. 1 4(a). 


6 44' W, 65 m, many colonies in spirit, coll. 'Atlantide' expedition, sta. 151, 16 Apr 1946, 
cf. 'L. pelagica\ det. W. Vervoort; RMNH, Leiden. Sete, S France, infertile material on three 
microslides, det. Clytia serrulata by Stechow, 1919a : 46; Munich Zoological Museum. 

DESCRIPTION OF HYDROID STAGE. Among the most variable of all Atlantic hydroids. 
Detached colonies can continue growing in the plankton and form one extreme of variation, 
while benthic colonies are more typical of the species and form the other. 

1. Benthic colonies. Colony comprising a creeping stolon from which short to long 
hydrothecal pedicels arise at irregular intervals. Hydrotheca thin walled, campanulate, 
length : breadth ratio 1-3, rim with 7-16 pointed to rounded or (?rarely) flat topped cusps, 
embayments typically pointed but sometimes rounded; hydrotheca usually round in cross- 
section but sometimes (Vervoort, 1968) sinuous near the top. Hydranth with the wide range 
of 20-30 tentacles (Hincks, 1868; cf. Obelia geniculata, p. 120). Pedicel straight, erect; 
annulated, usually with smooth central region and 2-10 annuli each end, rarely annulated 
throughout; sometimes branched, each branch having characteristic upward-curved basal 
region (similar to that in C. paulensis), branches often parallel with main pedicel; branch 
annulated basally and distally, central portion smooth. Gonotheca 3 = 9, broad, tubular, 
sometimes asymmetric, walls smooth to deeply concertinered, often with slight sub-terminal 
constriction; tapered below; aperture large, terminal; gonothecal pedicel short, on hydro- 
thecal pedicel or on stolon. Medusa released at four-tentacle stage. Nematocysts described by 

2. 'Planktonic' colonies. Floating colonies have been described by several authors (e.g. 
Van Breemen, 1905; Billard, 1907; Vervoort, \946a, 1959, 1968, 1972). Available evidence 
suggests that they are benthic colonies which have grown on sand and become detached (see 
Remarks). Colonies comprise one to several pedicels arising from a basal disc which often 
(Leloup, 1933) envelops a sand grain, stolons apparently being absent. Pedicels 
exceptionally long, branching repeatedly, branches often aligned parallel with primary 
pedicel. Hydrotheca long, narrow, thin walled, terminal cusps as in benthic form. Gonotheca 
long, smooth walled; rugose and concertinered gonothecae apparently not recorded from free 
floating material. 

Variation. Ralph (1957) found that cooler water increased the number of hydrothecal cusps. 
Although the species is highly variable there have been few studies of the factors involved. 

DISPERSIVE STAGE. A medusa, released at the four-tentacle stage. Umbrella hemispherical or 
slightly flatter; up to 25 mm in diameter; jelly fairly thin. Velum narrow; stomach short, 
four-sided; manubrium about 1/6 bell diameter in length, four-lipped. Radial canals 4; 
gonads elongate to oval, on distal 1/4-3/4 of radial canal but stopping just short of ring canal. 
Marginal tentacles 16-32 [58 recorded, Kramp, 1919 (=64)], short, hollow, smooth, with 
spherical bases. 1-3 closed marginal vesicles between each marginal tentacle and next. 
Tentacle bases and stomach yellow, yellowish brown, greenish or purple; gonads yellowish. 
(After Kramp, 1919, 1961; Russell, 1953.) 

The relation between hydroid and medusa has not always been understood, and there are 
still some unsolved problems relating to the nominal species described from the hydroid 
stage as 'Laomedea gracilis Sars'. 

Bohm (1878), working at Helgoland, worked out the life cycle of C. hemisphaerica and his 
account was quoted by Haeckel (1879 : 187) in his World synopsis; but many years later 
Mayer (1910 : 267) wrote 'the mature hydroid is not known with certainty, but is probably a 
Campanulina (see Hincks, 1868 : 179)'. Mayer should have quoted Hincks as indicating 
"Campanularia raridentata\ now regarded as conspecific with hemisphaerica. Hincks' 
citation in fact referred back to Wright's (1862) description of the hydroid stage under the 
name Thaumantias inconspicua; but Hincks (1852) himself had still earlier seen medusae 
released (identifying his material as "Campanularia volubilis Ellis & Solander'). Hincks thus 
seems to have been the first to record medusa release in the present species. Du Plessis (1871) 
also saw medusa release at an early date, but used the combination Clytia volubilis. (Notes on 


the widespread confusion between the species names volubilis and hemisphaerica are given 
below.) Mayer's oversight is the more surprising since he quoted MetschnikofTs (1886&) 
description of a polyp reared from the Mediterranean medusa Clytia flavidula (Peron & 
Lesueur, 1810a), which Mayer thought conspecific. The identity of C. flavidula is discussed 
under Remarks. 

The later but independent suggestions of Browne (1896 : 488; 1900 : 725-726; Browne & 
Vallentin, 1904:125, 127) that the hydroid of 'Phialidium hemisphaericum' was 
'Campanularia Johnston? were supported by Kramp in 1914 (Kramp, 1919 : 93, footnote), 
some forty years after Bohm and Haeckel wrote and even longer after the suggestions of Du 
Plessis, Hincks and Wright. The delay parallels that which occurred in the working out of the 
Obelia life cycle (Cornelius, \911a, b). 

REPRODUCTIVE SEASON. Medusae occur all through the year in British waters but are most 
numerous from spring to autumn [Hincks, 1852, as C. volubilis (in February); Russell, 1953; 
J. H. Robson, in Evans, 1978, as C. gracilis)]. Lo Bianco (1909, as Clytia flavidula and C. 
johnstoni) recorded medusae off Naples from August to October and from January to March. 

DISTRIBUTION. Nearly cosmopolitan in coastal waters (Ralph, 1957; Kramp, 1961), 
occurring in most of the present area. In European waters the medusa is one of the 
commonest (Russell, 1953). However, the species was said to be scarce in N Norway by 
Mathiesen (1928) who cited only two certain records, from Bergen and Hammerfest. It has 
been reported absent from Greenland (Kramp, 1943; Calder, 1970) although present in 
Iceland, Spitzbergen, the Faeroes and the Bering Sea (Calder, 1970). 'Planktonic' hydroid 
colonies have been reported as common in the southern North Sea and off much of the coast 
of W Africa (Vervoot, 19460, 1959). 

HABITAT. Usually benthic; intertidal to 150+ m (Crawshay, 1912; Marine Biological 
Association, 1957; Kramp, 1959; Millard, 1975; BMNH collections). Naumov (1969) 
reported a lower limit of 300 m but did not cite material. Rees & White (1966) listed a 
record from 1250 m off the Azores as C. gigantea, a species here regarded conspecific; but 
I have not checked this report. 

The species has been found on a wide variety of invertebrate and algal substrates, and there 
is no regular association. Among the unusual recorded substrates are sand grains (see 
Remarks) and pelagic cirripedes. There are several records on parasitic copepods 
themselves on fish (on Lernaeocera on Gadus by Leloup, 1930&; on Peniculus on Mullus, 
on Lernaeenicus on Clupea, and on Dinematura on Cetorhinus (the Basking Shark), all by 
Debouteville & Nunes, 1951, 1952). As in Obelia, long distance transport would seem to 
result (pp. 45, 120). 

REMARKS. Two factors have contributed to the profusion of redescriptions of this species 
and to the consequent number of synonyms: it is nearly cosmopolitan, and it is highly 
variable. The combination Clytia hemisphaerica was introduced only some fifteen years ago 
and there is still much use in a detailed analysis of the taxonomic history of the species. 
There has been confusion with other species and with other genera, and some of the 
problems have yet to be solved. The nominal species described from the NE Atlantic are 
considered below, so far as possible in chronological order. 

The notes by Stechow (192 la, \923a) and Rees & Thursfield (1965) on the nominal 
species Sertularia uniflora Pallas, 1766, were partly misleading. Pallas included in his 
synonymy Ellis' (1755) plate 14, figure A, Linnaeus' (1758) citing of that plate under the 
name Sertularia volubilis, and Baster's (1762) plate 2, figures 2a, b, d, 3, 4c, e, 7a-c. Ellis' 
illustration shows a campanulariid hydroid growing on Hydrallmania falcata (Linnaeus, 
1758). The pedicels were spirally grooved throughout, not annulated top and bottom as in 
the present species, and had a spherule at the upper end. These two characters indicate that 
Ellis' material was the species here called Campanularia volubilis. Hargitt (1909), Stechow 
and Rees & Thursfield wrongly supposed Ellis' material to be C. hemisphaerica. The name 
Sertularia uniflora Pallas, 1766, was in fact a nom. nov. for Sertularia volubilis Linnaeus, 


1758, and since both were based on Ellis' illustration uniflora is a junior objective synonym 

The other illustrations cited by Pallas under S. uniflora, those of Baster, show in one place 
(fig. 2A, b) a sharply cusped hydrothecal rim, but nowhere show details of the pedicel. 
Hence it cannot be determined whether Baster's material was volubilis or hemisphaerica. 
Gonothecae were not shown. Maitland (1876, quoted in Vervoort, 19460 : 276) referred part 
of Baster's illustrations (pi. 2, fig. 7A, a) to Orthopyxis Integra (Macgillivray, 1842), an 
opinion disagreed with by both Vervoort (19460) and myself as the hydrotheca of Integra 
does not have a cusped margin. Nevertheless, I here designate the material shown by Ellis 
(1755 : pi. 14, fig. A) lectotype of 5". uniflora Pallas, 1766, to provide formal protection for 
the name Integra from possible threat from uniflora should Maitland's opinion ever be 

Rees & Thursfield (1965) were further confusing in stating that 'earlier references by Ellis 
(17550, b) [sic] do not distinguish' between hemisphaerica and volubilis. They omitted the 
Ellis references from their paper: indeed, I cannot trace relevant papers by Ellis in that year. 
His only 1755 publication mentioning campanulariids was his book. In this Ellis (1755 : 25) 
stated clearly that there were two species and illustrated both on his plate 14. Linnaeus and 
Pallas (op. cit.) were each careful to cite only one of the species when referring Ellis' 
illustration respectively to their synonymies of volubilis and uniflora. The other illustrated 
species (Ellis, 1755 : pi. 14, B) was named Sertularia syringa by Linnaeus (1767 : 1311) and 
is now known as Calycella syringa, family Calycellidae (e.g. Cornelius, 1978). 

Thus there is little confusion in the early literature; but Rees & Thursfield (1965) were 
probably correct in stating that the 'hydroid [stage of C. hemisphaerica] is recognisable for 
the first time under the name Sertularia uniflora: Ellis, 1 768 : pi. 19, fig. 9'. And in this paper 
Ellis clearly did confuse the two species he had previously illustrated separately. His use of 
the name uniflora in that paper can be regarded as misidentification, not as homonymy. 
Ellis' (1768) illustrations show clearly the annulated, not spiralled, pedicels of hemi- 
sphaerica incidentally slightly unusual in lacking a smooth central portion and a 
characteristic 'concertinered' gonotheca. Further confusion might have resulted from Ellis & 
Solander's (1786) application of the name Sertularia volubilis to illustrations clearly of the 
present species, but the slightly earlier literature just discussed prevents it. However, Hargitt 
(1909) was not alone among later authors in applying the combination Clytia volubilis to the 
present species. 

C. hemisphaerica was placed in the genus Oceania Peron & Lesueur, 18100 (as O. 
flavidula), by those authors. 

Clytia urnigera Lamouroux, 1816, is referred to C. hemisphaerica but is discussed here 
under Orthopyxis Integra (pp. 64-65). 

The combination Laomedea gracilis Sars, 1850, has plagued the literature (see also 
Remarks under Gonothyraea loveni); but as noted by Vervoort (19460 : 285) the species 
name should not be used as it is preoccupied by "Lomedea gracilis'' C. Pickering, in Dana, 
1 846 : 689 (lapsus pro Laomedea gracilis), which I refer to Obelia dichotoma (p. 117). I 
propose the name Clytia sarsi nom. nov. in place of L. gracilis Sars, 1850 (preocc.). 
Lectotype material of L. gracilis Sars is designated here on page 94. Nevertheless, the name 
gracilis has been widely applied to colonies having long, narrow hydrothecae and smooth 
gonothecae. But following Ralph's (1957) account of variation in New Zealand populations 
most authors have regarded '50r$/-type' colonies as an extreme variation of hemisphaerica. 
Further evidence was provided by W. J. Rees (in Rees & Thursfield, 1965) who reared a 
medusa from a sarsi ('gracilis'') colony and found it the same as the medusa hemisphaerica s. 
str. Kiihl (1967) regarded 'sarsi-type' colonies conspecific with those of hemisphaerica on 
more subjective grounds. But Ostman (1979) separated the species on the fine structure of the 
nematocysts (see also p. 42) and the problem is not yet resolved. 

There has been some further debate over the degree of affinity between C. sarsi (=L. 
gracilis Sars, 1850), and Campanularia pelagica Van Breemen, 1905. Some authors have 
regarded them identical (e.g. Leloup, 1933, 1952; Vervoort, 19460; Naumov, 1960, 1969; 


Garcia Corrales et al., 1978); but the validity of C. pelagica s. str. has been difficult to assess. 
(Campanularia attenuata Calkins, 1899, from Puget Sound, seems similar.) Among recent 
authors Vervoort's (1946a, 1959, 1968, 1972) descriptions of the pelagica 'form' have been 
important but the possibility has remained open that pelagica was simply based on sarsi or 
hemisphaerica material which had detached and grown in the plankton. Billard (1917) also 
had failed to resolve the question but E. T. Browne, A. Kemna and E. Leloup (in Leloup, 
1933) thought sarsi (gracilis) just to be detached, floating colonies of hemisphaerica. 

Vervoort (1959, as Laomedea pelagica) reported that 'floating' colonies occurred in the 
North Sea and along much of the west coast of Africa, but Millard (1966) referred Vervoort's 
African coast material to C. hemisphaerica without comment. Subsequently Vervoort (1968) 
reaffirmed his view that pelagica was distinct. He had not seen 'intermediate material'; but 
presumably there is no intermediate environment. Vervoort separated pelagica on its 
slender, gradually narrowing hydrothecae, an undulating cross-section just below the 
hydrothecal rim and a smooth walled gonotheca. He cautiously noted that two species of 
Clytia medusae had been recorded from the North Sea by Russell (1953) and that these 
might relate to the two hydroids, pelagica and hemisphaerica. He subsequently (Vervoort, 
1972) retained specific status for pelagica but Millard (1975) again referred Vervoort's 
African coast material to hemisphaerica. So the experts were divided. Naumov (1960, 1969) 
meanwhile had considered pelagica distinct but extended its limits slightly further than 
Vervoort to include sarsi-lype material with smooth walled gonotheca. Garcia Corrales et al. 
(1978) similarly regarded sarsi + pelagica distinct from hemisphaerica, based on sharper and 
longer hydrothecal cusps and longer hydrotheca in the former; and like Vervoort (1968) had 
seen no intermediate material. Other features on which pelagica has been distinguished are 
the frequency of branching and the tendency of the branches to run parallel with the primary 
stem, noted by Vervoort (1959) but not apparent in all planktonic material; and a smooth, 
asymmetric gonotheca (e.g. Billard, 1917). 

Vervoort (1972 : 91, footnote) drew attention to the report by Sars (1850, 1857; ?repeated 
in Leloup, 1952) that 'pelagica' had sessile gonophores. Vervoort cited observations that 
'pelagica-type' material released its medusae and (pers. comm.) now considers that the 
contrary observations of Sars might have been made on Gonothyraea loveni material. 

Old and new evidence for uniting pelagica with hemisphaerica runs as follows. Ralph 
(1957) showed that smooth gonothecae are not unusual in hemisphaerica, and that the 
length of the hydrotheca and shape of the cusps on the rim vary widely, to an extent which 
encompasses the three nominal taxa. The basal discs (=Haftplatte of Kuhn, 1913, and 
disque de fixation of Houvenaghel-Crevecoeur, 1973; there seems no English equivalent 
already) of attached C. hemisphaerica examined by me seem identical with the 'partie 
basale' or "pied . . . globuleuse' described in planktonic colonies by Billard (1917) and Leloup 
(1933) under the name pelagica (Fig. 9). Also, Leloup showed that the basal discs of these 
free-floating colonies often contain a sand grain, indicating a benthic origin. Next, the 
distinctive upward-swept pedicel bases in hemisphaerica s. str. seem identical with those 
described as pelagica. Finally, it might have been expected that a hydroid which was 
habitually planktonic would have some obvious modification to that end; but there seems 

The relation between the three nominal forms hemisphaerica, sarsi and pelagica is 
certainly close, and the relation tojohnstoni is still uncertain (see below). Whether the differ- 
ences are phenotypic or genotypic cannot yet be decided; but on skeletal and medusa 
characters it seems best now to interpret the variation as representing a single 'morphological 
species'. More detailed studies, such as that of Ostman (1979) on the nematocysts, might 
shed further light. 

Campanularia volubilis (Ellis & Solander, 1786) sens. Hincks (1852) was the present 
species. Hincks' account was probably the earliest record of medusa release in C. 

Campanularia johnstoni Alder, 1856a, is apparently the earliest available name for the 
hydroid stage most usually (but subjectively) connected with the medusa on which the 


present species is based (see Nomenclature, above). It is also the correct species name for the 
type species of the genus Clytia (p. 71). The type series of Campanularia johnstoni is 
mentioned under Material, above, and by Cornelius & Garfath (1980). See also Cornelius 

Phialidium viridicans Leuckart, 1 856, is discussed above (p. 7 1 ) and on page 86. 

Clytia noliformis (McCrady, 1857, as Campanularia) was founded on a hydroid which 
released a Clytia medusa lacking gonads when liberated. Kramp (1959, 1961) regarded the 
lack of gonads on release an important character; and the species has been widely recognized 
from both hydroid and medusa stages in warm waters throughout the World (references in 
Fraser, 1944; Kramp, 1961; comment in Rees & Thursfield, 1965). Mammen (1965) gave 
this name to a medusa he reared but it closely resembled Russell's (1953 : pi. 17, fig. 6) 
illustration of C. hemisphaerica. Mammen's medusa differed only in not showing gonads till 
48 hrs old. C. noliformis has not otherwise been redescribed since Kramp (1961) wrote and 
may prove conspecific. Picard (1949) referred medusa and hydroid material from Ville- 
franche to noliformis but his specimens, like Mammen's, seemed to differ from hemi- 
sphaerica only in the short delay in acquiring gonads. Later he included the species in a 
Mediterranean faunal list, regarding C. jlavidula Metschnikoff, 1886a, and C. mollis 
Stechow, 1919a, as conspecific (Picard, 19586). These two species have both been referred 
to hemisphaerica by at least some senior authors; and are discussed in their chronological 
place below. Rees & Thursfield (1965) referred some Cape Verde material from James 
Ritchie's collection to C. noliformis. They regarded Campanularia ptychocyathus Allman, 
1888, from the Azores, as conspecific; but Stechow (1925 : 521) treated ptychocyathus as 

Campanularia gegenbauri Sars, 1857, was based on a fertile hydroid colony illustrated 
by Gegenbaur (1854: pi. 1, figs 1, la, as Campanularia n. sp.). I concur with Hincks 
(1868 : 145), Bohm (1878 : 168) and Bedot (1910 : 254) that the species should be referred 
to Clytia hemisphaerica. 

Campanularia volubiliformis Sars, 1857, was a name applied to material described as 
Campanularia sp. by Gegenbaur (1854). The shape of the hydrotheca, pedicel and 
gonotheca illustrated indicate the present species and not Campanularia hincksii as 
suggested by Carus (1844) and Broch (19126). Heller (1868) and Picard (195 la) reported 
further material without description; while Stechow 1919#:70) referred the species to 

Campanularia raridentata Alder, in Hincks, 18616, was referred to Clytia hemisphaerica 
by Billard (1928) and Rees & Thursfield (1965), and provisionally by Vervoort (1968). The 
synonymy was agreed by Cornelius & Garfath (1980), who alone saw the holotype. The 
specimen was simply a young colony of C. hemisphaerica. Rees & Thursfield wrongly 
ascribed the original description to 'Alder, 1857'. 

Clytia bicophora Agassiz, 1862, originally based on hydroid material from New England, 
was recorded as a medusa in the Gulf of Trieste by Thiel (1935 : 172, as Phialidium 
bicophorum). But Agassiz' detailed description and clear illustrations appear to represent C. 
hemisphaerica, and bicophora can be regarded conspecific. Kramp (1959 : 149) listed other 
records of bicophora as 'uncertain', and indeed Bohm (1878) referred bicophora to C. 
johnstoni, which is also regarded conspecific. Kramp (1959, 1961) nevertheless described a 
medusa under the name bicophora and listed references to that species; but the supposed 
differences from hemisphaerica seem unimportant. 

Clytia (Platypyxis) cylindrica Agassiz, 1862, was based on both hydroid and medusa 
material from Massachusetts Bay and Buzzards Bay, North America. The species was 
referred to C. hemisphaerica by Bohm (1 878), but to C. noliformis (McCrady, 1 857) by Bedot 
(1910:348) and Kramp (1961 : 170). (Bedot clearly regarded Platypyxis a synonym of 
Clytia, and I concur; see p. 71.) Agassiz' description of cylindrica includes no important 
differences from hemisphaerica and like Bohm I regard them conspecific. Vervoort 
(1968) described new material, and commented on the similarity of C. elsaeoswaldae 
Stechow, 19 14. 1 agree with Vervoort in regarding the Stechow species conspecific. 


The combination Phialidium languidum (Agassiz, 1862) has been applied to medusae 
caught off Senegal, but Kramp (1955) referred both the material involved and the nominal 
species itself to C. hemisphaerica (see Kramp, 1933, 1955, 1 96 1 , for discussion). 

Campanularia ?gigantea Hincks, 1866, was probably based on large hemisphaerica 
hydroid material. The eleven hydrothecae of the holotype well enough preserved for study 
measured, in mm from diaphragm to tips of hydrothecal cusps, M8, 1'26, l - 29 (3), 1'53, 
1-62 (2), 1*78, 1'88, 1-91. The wide range is striking, as is the sheer size of the largest; but 
Hincks was rightly cautious to introduce the name as ?gigantea. Several authors have 
referred large material to gigantea solely on the basis of size. The pedicels too are long, but 
the upward curve of the pedicel bases recalls hemisphaerica s. str. and continued separation 
seems uncalled for. Gonothecae have never been reported in this large material. Hincks' 
(1868) later illustration of the holotype is accurate; and that of Jaderholm (1909), of 
non-type material, is similar. The material fron N Scotland listed by Rees & Thursfield 
(1965; BMNH 1964.8.7.75) is C. hemisphaerica and is smaller than the holotype of 
?gigantea. The material from Belgium cited by Leloup (1952) seems to have been the same, 
as does the North Sea material listed by Billard (1928, as Clytia johnstoni). 

Gastroblasta raffaelei Lang, 1886, was based on a medusa from Naples. The species was 
provisionally referred to C. hemisphaerica by Kramp (1959 : 148). 

Clytia flavidula: Metschnikoff, 1886<2, was a comb. nov. for Oceania flavidula Peron & 
Lesueur, 1810a, originally described from medusae collected from Nice. Mayer (1910) 
referred the species to C. hemisphaerica and Russell (1953) concurred. Lo Bianco 
(1909 : 540) also used the combination Clytia flavidula. Kramp (1961 : 65) listed another use 
of the binominal O. flavidula, for O. armata, a clavid medusa. 

The Mediterranean material referred by Stechow (191 9a) to Campanularia serrulata Bale, 
1888 (first described from Sydney), was examined here and found to be simply Clytia 
hemisphaerica; as indeed seems Bale's species, which was described from infertile material. 
In general agreement, Stepanyants (1979) referred C. serrulata to 'Campanularia gracilis' 
Sars, 1850, itself regarded conspecific (see above). Picard (1958/7) recorded the species as 
Mediterranean without comment. 

Campanularia attenuata Calkins, 1899, originally described from Puget Sound, resembles 
'Laomedea pelagica Van Breemen, 1905', and like it should probably be regarded 
conspecific with Clytia hemisphaerica. Material was recorded from Ghana by Buchanan 
(1957) and it is well known that 'pelagica-type' colonies occur along the African coast (see 
above). Although West & Renshaw (1970) recognized the species on the basis of its branched 
colonies as grown in vitro at certain temperatures (see p. 40) further proof of validity seems 
necessary. C. attenuata Stechow, \9\9a, from Villefranche, is a junior homonym and later in 
the same paper Stechow (1919a:157) introduced the nom. nov. Campanularia 
villafrancensis instead. However, villafrancensis too seems conspecific with hemisphaerica. 

Campanularia edwardsi Nutting, 190 \b, based on material from Massachusetts, was 
redescribed at length by Kubota (1978, as Clytia). He provisionally referred C. edwardsi to 
the present species, and I concur. 

Clytia simplex Congdon, 1907:471-472, figs 14-15, based on material taken off 
Sargassum weed near Bermuda, was reported south of the Azores by Vanhoffen (1910). 
Fraser (1944), however, referred the species to C. noliformis McCrady, 1857, here regarded 
conspecific with C. hemisphaerica; and it seems probable that C. simplex too is conspecific. 

Clytia obeliformis Stechow, 1914, was based on material from Bergen, Norway. The 
original illustration shows both hydrotheca and gonotheca typical of C. hemisphaerica and 
the type material, examined here, comprises simply a rather elongate colony of the present 
species. There is no reason to maintain a separation. 

Campanularia acuta Stechow, 1919a, was a nom. nov. applied to C. raridentata: var. 
Marktanner-Turneretscher, 1890, based on material from NW France. The variety was 
originally held to differ from C. raridentata s. str. in lacking annulations around the base of 
the gonotheca; and like that species seems to have been based on Clytia hemisphaerica 



Campanularia brachycaulis Stechow, 1919a, was based on a small, infertile colony from 
Villefranche. The illustrated hydrotheca does not seem to differ from that of Clytia 
hemisphaerica; and I must disagree with Patriti (1970) who referred the species to 
Campanularia hincksii. 

Clytia compressa Totton, 1930, was proposed on the basis of the shape of the hydrothecal 
cusps and of the gonotheca. Ralph (1957) and Vervoort (1968) regarded it conspecific, and I 

Stolon growth in the hydroid stage was described by Hale (1964, \913a, b). 

Clytia hummelincki (Leloup, 1935) 
(Fig. 10) 

Laomeda hummelincki Leloup, 1935 : 19, fig. 7; Buchanan, 1957 : 360, fig. 1 1 a-b. 
Campanularia hummelincki: Fraser, 1944 : 122, pi. 21, fig. 93; Vervoort, 1966 : 96. 
Clytia hummelincki: Deevey, 1954 : 270; Millard, 1966 : 480-481, fig. 14g-l; Millard, 1975 : 218-220, 
fig. 72 f-h. 

TYPE LOCALITY AND MATERIAL. Isle de Bonaire, West Indies, 0'7 m depth, on dead coral, 
infertile material on microslide (Fig. 10); Mus. Roy. Hist. Nat. Belg. collection. 

OTHER MATERIAL EXAMINED. Fragments of fertile colonies on two microslides, Aghulas Bank, 
off South Africa (34 43' S, 25 40' E); South African Museum reg. no. H 2967 (mentioned, 
Millard, 1966, 1975). 

DESCRIPTION OF HYDROID STAGE (partly after Leloup, 1935; Buchanan, 1957; Millard, 
1975). Colony a tortuous stolon bearing long hydrothecal pedicels at irregular intervals; 

Fig. 10 Clytia hummelincki. (a-b) hydrotheca from syntype series, two magnifications, (c) 9 
gonotheca (after Millard, 1975 : fig. 72H). Scales: (a) 50 /zm; (b-c) 500 //m. 


gonothecae subsessile on stolon. Hydrothecae short, wide, unthickened, length = maximum 
breadth, tapering sharply towards base which is usually slightly constricted; rim often 
sloping, even to slightly and irregularly sinuous; diaphragm delicate, oblique in lateral view, 
either parallel with hydrothecal aperture or sloping in opposite direction; basal chamber 
small. Hydrothecal pedicel long, smooth or with several basal annulations and up to c. five 
groups of 3-5 annuli along length; sub-hydrothecal spherule present, below which often a 
slight constriction (see Fig. 10). Hydranth with spherical hypostome and c. 20 tentacles. 
Gonotheca (?rf = 9) sessile to shortly stalked, truncate; widest above, tapering below; 
sometimes asymmetric; aperture probably broad as end; with 1-2 medusa buds, develop- 
ment successive; buds with 4 radial canals and 4 tentacle rudiments (South African material); 
mature gonotheca and medusa undescribed. 


Table 2 Clytia hummelincki Measurements in /zm. 

Caribbean Ghana South Africa 

(Leloup, 1935) (Buchanan, 1957) (Millard, 1966, 1975) 


Length 200-240 250 250-^00 

Breadth (max) 250 240-250 200-420 

Length/breadth 0-8-0-96 1-00-1-04 0-7-1-36 


Length 1500-2000 2000 1920^730 

Breadth (max) 40-60 90-150 


DISPERSIVE STAGE. Probably a medusa. Millard (1966, 1975) noted medusa buds with 4 
marginal bulbs and so referred the species to Clytia. See also Description, Remarks under 
C. hemisphaerica and Dispersive stage under C. gravieri. 

REPRODUCTIVE SEASON. The only recorded fertile material was collected from Aghulas 
Bank, South Africa, on 10 February, 1962 (Millard, 1966). The gonothecae contained well 
developed medusae. 

DISTRIBUTION. A little known species, recorded in the NE Atlantic only from Ghana 
(Buchanan, 1957). Other Atlantic records indicate a wide distribution: West Indies (Leloup, 
1935; Vervoort, 1966), Florida Keys and Woods Hole (Deevey, 1954, possibly northernmost 
record of species) and South Africa (Millard, 1966, 1975). 

HABITAT. The few collected colonies have been on Lepas (Cirripedia) attached to a buoy, on 
sublittoral coral debris and on intertidal Sargassum weed (Leloup, 1935; Buchanan, 1957; 
Millard, 1966), indicating a wide substrate range. Recorded depths have so far ranged only 
from intertidal (Buchanan) to less than 1 m (Millard). 

REMARKS. This rarely reported species is clearly widespread in the warmer parts of the 
Atlantic Ocean. It is retained in Clytia following Millard's notes on the developing medusa. 
Although the species has a sub-hydrothecal spherule it is not transferred to Campanularia 
since that character might be due to convergence (see p. 41). But the affinities of the species 
remain uncertain and the identity of the medusa might provide better indication. 



Clytia islandica (Kramp, 1919) 
(Fig. 11) 

Phialidium islandicum Kramp, 1919 : 95, pi. 4, figs 1 1-13, pi. 5, figs 1-2; Russell, 1953 : 294-296, 
text-figs 180-181; Kramp, 1959: 149, 215, 218, 221, 222, fig. 190; Kramp, 1961 : 169 (?syn. 
Staurostoma laciniatum var. hybridum Le Danois). 

TYPE LOCALITY. Coastal waters of Iceland. 

DESCRIPTION OF MEDUSA STAGE (Hydroid unknown). Diameter of adult 35-40 mm, 
umbrellar saucer-shaped, jelly thin; stomach small, cruciform; manubrium reduced, mouth 
square surrounded by crenulated lips; gonads narrow, along almost whole length of radial 
canals; tentacles c. 200, alternating with statocysts. 
Variation. See Russell ( 1 953) and Kramp ( 1 959). 

Fig. 1 1 Clytia islandica. Redrawn after Kramp ( 1 959 : fig. 1 90). Diameter 35-40 mm. 

DISTRIBUTION. Reported at least as far south as SW Ireland but records generally more 
northerly (Russell, 1953). Kramp ( 1 96 1 ) cited unconfirmed records from the Bay of Biscay. 

REMARKS. Distinguished from the medusa stage of Clytia hemisphaerica by: larger diameter 
(up to 40 mm, not up to 20 mm), flat (not hemispherical) shape, more numerous tentacles 
[up to c. 200, not up to (rarely) 58], usually rather larger gonads and one (not 1-3, usually 2) 
statocysts between tentacles. The hydroid is not known. 

Disuse of the genus name Phialidium is discussed above (p. 74). Kramp (1961) referred 
the genus Staurostoma Haeckel, 1879:130, to Staurophora Brandt, in the family 

Clytia linearis (Thornely, 1 899) 
(Fig. 12) 

Obelia linearis Thornely, 1899 : 453, pi. 44, fig. 6. 

Campanularia gravieri Billard, 1904a : 482, fig. 1; Billard, 1907 : 171-172. 
?Clytia geniculata Thornely, 1904 : 1 12-1 13, pi. 3, figs 4, 4a. 
Campanularia ?obHqua Clarke, 1907 : 9, pi. 5, figs 14. 

Clytia linearis: Stechow, 1913 : 66-69, figs 23-25; Hirohito, 1977 : 14-20, fig. 4a-j (syn. Campanularia 
gravieri Billard; Clytia hendersonae Torrey, 1904; C. alternata Hargitt; Laomedea bistriata Leloup). 
Clytia alternata Hargitt, 1924 : 483, pi. 2, fig. 7. 
tClytia (?)foxi Billard, 1926 : 93-94, fig. 9A-B. 



Laomedea (Obelia) bistriata Leloup, 193 la : 4, figs 8-1 1 . 

Clytia gravieri: Billard, 1938 : 429-432, figs 1-3, ?fig. 4 (syn. C. alternata Hargitt, Laomedea bistriata 

Leloup); Picard, 1955 : 185-186; Millard & Bouillon, 1973 : 51-54, fig. 7e-g (syn. Obelia striata 

Clarke, 1907; C. serrata Millard, 1958); Millard, 1975 : 215-217, fig. 71 F-H (syn. Obelia striata 

Clarke, 1907); Garcia Corrales, Inchaurbe & Mora, 1978 : 29-30, fig. 12. 
Clytia obliqua: Picard, 1950 : 51-52. 
Campanularia (Clytia) gravieri: Vervoort, 1967:50-52, fig. 16 (syn. Clytia alternata Hargitt; 

Laomedea bistriata Leloup). 

TYPE LOCALITY. Blanche Bay, New Britain, Bismarck Archipelago. Material not located. 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Campanularia obliqua Clarke, 1907, infertile 
colonies on sertularian hydroid, in spirit, 'Perico Island', Gulf of Panamar, coll. r.v. 
'Albatross'; Smithsonian Institution cat. no. 29616. 

Clytia alternata Hargitt, 1924, fertile colonies preserved on microslide, Port Galero, 
Mindoro, Philippines; Smithsonian Institution cat. no. 42644 (Fig. 12). 


DESCRIPTION OF HYDROID STAGE (partly after Thornely, 1899; Billard, 1904<a, 1938; 
Stechow, 1925; Picard, 195 la; Vervoort, 1967; Millard & Bouillon, 1973; Millard, 1975; 
Hirohito, 1977). Colonies both stolonal and erect. When erect branching sympodially, up to 
c. 10 hydrothecae in extent, to 20 mm. Hydrothecal pedicels finely ringed throughout (6-37, 
usually c. 20) or with smooth central portions; distal pedicels shorter than proximal. 
Internodes of erect stems arcuate, narrowly ringed basally. Hydrothecae long, sides parallel 
to slightly divergent, often with slight asymmetric bulge; with 8-16 long narrow marginal 
cusps recalling those of Clytia paulensis and Obelia bidentata but each with internal 

Fig. 12 Clytia linearis. The specimen illustrated is a syntype of C. alternata Hargitt, 1924, here 
regarded conspecific. (a) part of colony, (b-c) hydrotheca and part of rim. (d) $ gonotheca. 
Scales: (a) 500 /mi; (b) 250 /mi; (c-d) 500 //m. 


stiffening strip reaching to tip and extending downwards sometimes to middle of hydrotheca; 
intervening bays rather wide and square, bulging out; diaphragm usually oblique but some- 
times transverse. Hydranth with 12-15 tentacles (in holotype specimen of C. alternata 
Hargitt). Mature gonotheca (?rf = 9) elongate, on 2-3 ringed pedicel, either on stolon or on 
erect shoot in axil; widest in centre, tapering below and usually above; one-flapped circular 
deciduous operculum; young gonotheca shorter, more sharply truncated. Gonophore with 
one or more rows of developing medusae; pre-release medusae with hemispherical umbrella 
and four marginal tentacles. 

Variation Hydrothecal length varies greatly, at least from 400 /^m to 1100//m (Millard, 
1975). Internodes and pedicels are shorter in upper parts of the colony (Hirohito, 1977). 
Hirohito found little difference between Red Sea and Japanese specimens, indicating a rather 
constant phenotype. Hydrothecal pedicels were on the whole shorter in the Japanese 
specimens, but gonothecae were similar in size. Billard (1938) reported that the embayments 
between the hydrothecal cusps are often distorted in preserved material due to lateral 

DISPERSIVE STAGE. A medusa, with hemispherical umbrella and four marginal tentacles on 
release; earliest stage identical with newly liberated Clytia hemisphaerica medusa, even in 
cnidom according to Picard (195 la). Adult medusa not described (Millard, 1975). Russell 
(1953) noted that the Mediterranean medusa Phialidium viridicans Leuckart, 1856, might 
prove valid. It might follow that Clytia linearis is its hydroid, but evidence is only 
circumstantial and the hydroid stages of C. hummelincki and C. paulensis must also be 
considered. 'P. viridicans' is here provisionally referred to C. hemisphaerica. 

DISTRIBUTION. Circumglobal in tropical to warm shallow seas (Millard & Bouillon, 1973; 
Hirohito, 1977) extending northwards in the Atlantic Ocean at least to the south coasts of 
France ('extremely common' intertidally at Banyuls, Picard, 195 la) and Spain (10-20 m off 
Alicante, Garcia Corrales et al., 1978). Billard (1907) recorded the species south of Madeira; 
Picard (1955) from Algeria; Rees & Thursfield (1965, as C. striata) from the Cape Verde Is 
and Rees & White (1966, as C. striata) from the Azores. Vervoort (1967) gave a list of known 
Indo-Pacific localities; and some possible records under other specific names are discussed 

HABITAT. On cirripede and pteropod shells and on other hydroids (Billard, 1904a; Vervoort, 
1967; Millard & Bouillon, 1973; Millard, 1975; Hirohito, 1977); also on intertidal rocks 
(Picard, 195 la). Intertidal (Picard) to 110m (Billard, 1907; Millard). 

REMARKS. Ritchie (1907) recorded material from the Azores as Clytia geniculata Thornely, 
1904, a name originally applied to specimens from Sri Lanka. Rees & Thursfield (1965) 
tentatively thought C. geniculata conspecific with C. striata (Clarke, 1907), which Millard & 
Bouillon (1973) and Millard (1975) referred to the present species. But Hirohito (1977) 
described what he considered undoubted C. striata material (on a pteropod) and considered 
the species valid. 

Picard (1950) assigned material from near Marseille to Clytia ?obliqua Clarke, 1907) but 
that species seems identical with Clytia linearis. Clarke based the distinction merely on the 
angle of slope of the hydrothecal cusps. The type material of C. obliqua, examined here, does 
not otherwise differ from the original description of C. gravieri. 

The type material of Clytia alternata Hargitt, 1924, was examined also by Hirohito 
(1977). As he stated, it resembles the description of C. linearis so closely that the two species 
can be regarded conspecific. C. foxi Billard, 1926, was based on slight differences and may 
also be the same species. Vervoort (1967) drew attention to the close similarity between C. 
alternata and C. gravieri Billard, 1904, which also seems conspecific. Billard (1938) had 
earlier thought them conspecific along with Laomedea bistriata Leloup, 193 la. He saw four 
marginal tentacle bulbs in the pre-release medusa and hence assigned the species to Clytia. 
Detailed discussion of several Pacific forms was provided by Hirohito. 


Clytia mccradyi (Brooks, 18886) 
(Fig. 13) 

Oceania sp. Brooks, 1888a : 29-30. 

Epenthesis mccradyi Brooks, 18886 : 147-1 62, pis 13-15; Sigerfoos, 1893 : 106. 

Oceania mccradyi: Mayer, 1900 : 50, pi. 2 1 , figs 56-59. 

Phialidium mccradyi: Mayer, 1910 : 271-272, pi. 34, figs 2-3, pi. 35, figs 1-3; Kramp, 1959 : 149; 

Kramp, 1961 : 170; Bougis, 1963 : 2,2-2,3, pi. 9, fig. 2. 
Further references were given by Mayer (1910) and Kramp ( 1 96 1 ). 

TYPE MATERIAL AND LOCALITY. Green Turtle, 1886, and Nassau, 1887, Bahamas (medusa 
stage only); material not located. 

MATERIAL EXAMINED. None available. 

DESCRIPTION OF HYDROID STAGE. The identity of the hydroid stage is unclear. The species 
was originally based on mature medusae collected from the plankton. Brooks (18886) later 
saw young medusae released from a hydroid colony and, identifying the young medusae as 
the same species as the adults earlier described, concluded he had found the hydroid. Mayer 
(1910) was sceptical. Brooks' description of the hydroid did not differ from the hydroid of C. 
hemisphaerica. The description included stolonal gonothecae with lateral constrictions 
characteristic of the better known hemisphaerica. Only Brooks has reported a conventional 
hydroid stage. Others (Sigerfoos, 1893; Mayer, 1910; Bougis, 1963) have described a much 
reduced hydroid stage comprising only hydroid blastostyles within gonothecae, attached to 
the gonad of the medusa. 

Fig. 13 Clytia mccradyi. Redrawn after Mayer (1910 : pi. 35, fig. 1 ). Diameter c. 15mm. 

DISPERSIVE STAGE. A medusa. The following description is after Brooks (18886) and Mayer 
(1910). Bell shallow, c. 15 mm diameter, less than half as high as broad, flexible; stomach 
short, less than l/8th as long as diameter of bell, quadrate, with four simple, slightly recurved 
lips; gonads short, oval, about mid-way between axis and bell margin; velum well developed; 
1 6-24 long, contractile marginal tentacles (type material had 1 6 only), 'many times as long 
as diameter of bell', each with statocyst at base; 8-16 other marginal statocysts; 1-4 (?4+) 
gonothecae borne on one to all of the gonads in many individuals. Gonads of medusa 
develop either eggs or sperm, or into hydroid blastostyles which produce medusae direct 
(details in Sigerfoos, 1893 and Mayer, 1910). 4-8 tentacles on release (Brooks, 18886). 

REPRODUCTIVE SEASON. Brooks (18886) recorded fertile hydroid material in June, 1887, at 
Nassau, Bahamas; Mayer (1910) in July at Tortugas, Florida. Medusae from April to July 

DISTRIBUTION. In the eastern North Atlantic I know of only one record: Villefranche, S 
France, found 'from time to time' (Bougis, 1963). Reported elsewhere in the World from 
the Bahamas and Florida (Kramp, 1959, 1961). 


HABITAT. Brooks (18886) reported the 'free' hydroid stage on algae in the harbours of Nassau 
and Green Turtle, Bahamas. 

The hydroid stage is at least sometimes partially suppressed and attaches to or grows on 
the medusa. Thus the species seems adapted to an oceanic existence and might be found 
away from coastal waters. Possibly the isolated record from the Mediterranean is evidence of 
a natural trans-atlantic dispersal. 

REMARKS. The unusual life cycle of this species has been checked by at least three 
independent workers (Sigerfoos, 1893; Mayer, 1910; Bougis, 1963) since the original descrip- 
tion by Brooks (18886). Brooks alone claimed to have found a benthic hydroid stage, but 
Mayer evidently doubted his observation. Although Mayer studied live medusae of the 
species closely, Brooks' account is explicit and suggests strongly that the benthic hydroids he 
described were of this species. At the time Mayer wrote such plasticity in the life cycle of a 
single hydromedusa species would have been regarded unusual and requiring more rigorous 
proof than today, when several such examples are known (summary in Naumov, 1969). 

Brooks stated that the medusa has either 4 or 8 tentacles on release, a departure from the 
unvarying four usually considered normal in this genus (see p. 42). 

Clytia paulensis (Vanhoffen, 1910) 
(Fig. 14) 

Campa nularia pa ulensis Vanhoffen, 1910 : 298, fig. 19. 

Clytia ulvae Stechow, 19190 : 47-48, fig. N. 

Clytia paulensis: Stechow, 19230: 110, fig. N; Philbert, 19356:25-26, fig. 4; Picard, 1955: 186; 

Millard, 1966 : 481-483, fig. 15 (?syn. C. ulvae Stechow, 1919a); Millard, 1975 : 221, fig. 73a-d. 
Obelia paulensis: Naumov & Stepanyants, 1972 : 37, fig. 2a-b. 

TYPE LOCALITY AND MATERIAL. Shallow water in crater lagoon of St Paul Island, S Indian 
Ocean (38 40' S, 77 34' E), 26 Apr 1903, on Sertularella polyzonias (Linneaus, 1758), coll. 
Deutsche Siidpolar-Expedition 1901-1903; material not located. 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Clytia ulvae Stechow, 1919<2, infertile colony 
on Ulva (green alga), Marseille, microslide preparation; Munich Zoological Museum. Also 1 
non-type specimen, 'Valdivia' sta. 100 (S Africa), det. E. Stechow as C. ulvae; MZM. 

OTHER MATERIAL EXAMINED (All BMNH material is listed. None is fertile.). Breakwater at S 
end of Landguard Pt, Felixstowe, Suffolk, SE England, ELWS, Sep-Oct 1976, infertile 
colony on Crisia sp. (Bryozoa) itself on Tubularia indivisa Linnaeus, spirit + 1 microslide 
preparation, coll. & det. R. G. Hughes; 1977.5.7.2. Off Berry Head, Tor Bay, Devon, SW 
England, c. 15m, Mar-Jun 1973, infertile colony on Nemertesia sp., spirit* 1 microslide 
preparation, coll. R. G. Hughes; 1973.8.13.1 (see Remarks; mentioned, Hughes, 
1975 : 291). Off Berry Head, c. 15m, summer 1974, infertile colony on Nemertesia sp., co- 
epizoic with C. hemisphaerica, spirit + 1 microslide preparation, coll. & det. R. G. Hughes; 
1977.5.7.1 (Fig. 14). Mewstone Ground, near Plymouth, Devon, Oct 1899, infertile colony 
on Laomedea flexuosa, 1 microslide preparation, coll. Marine Biological Association of 
U.K., ex E. T. Browne colln; 1961.11.14.16 (see Remarks). R. Ranee, nr St Malo, NW 
France, infertile colony on Hydrallmania falcata (Linnaeus, 1758), spirit + 2 microslide 
preparations, coll. M. Philbert; 1935.7.10.1 (?mentioned, Philbert, 19356). Sta. SCD258 W, 
Univ. Cape Town Ecol. Survey, 14 Jul 1961, infertile colonies on Obelia dichotoma, spirit + 
1 microslide preparation, pres. N. A. H. Millard; 1962.6.18.9. 

DESCRIPTION OF HYDROID STAGE. Colony stoloniferous, comprising a tortuous, branched, 
mostly unringed stolon from which usually unbranched pedicels arise at intervals. Pedicels 
long, occasionally branching as in C. hemisphaerica with similar upward-curved bases 
bringing branches approximately parallel with main pedicel; annulated basally, below 
hydrotheca and sometimes centrally, forming 1-2 smooth central portions. Hydrotheca 



Fig. 14 Clytia paulensis. (a) single hydrotheca on pedicel (right) adjacent to same of C. 
hemisphaerica (left), SW England; 1977.5.7.1. (b) part of (a), right hand specimen, enlarged, (c) 
gonotheca (after Millard, 1975 : fig. 73D). Scales: (a, c) 500 /mi; (b) 100/zm. 

rather longer than in C. hemisphaerica, length : breadth ratio usually 3-4, exceptionally 1^; 
7-1 1 rounded bimucronate cusps; flared appearance immediately below rim resulting from 
outward bulging of larger embayments; diaphragm oblique; longitudinal folds in hydrothecal 
wall in microslide preparations can look like striations (Millard, 1975) but are artefacts. 
Hydranths in present material with 16-22 tentacles. Gonotheca (not seen) ?d" = 9, cylindrical, 
tapering gradually below and slightly above, smooth (after Stechow, 19230; Philbert, 19356; 
Millard, 1975); borne on stolon on short annulated pedicel. 1-3 medusa buds per blastostyle, 
pre-release medusa with 4 tentacle buds (Millard, 1975); free medusa not yet described. 
Variation. Cusps on hydrothecal rim variable in both length and breadth; always rounded in 
present material. Embayments between cusps irregular in depth so that adjacent ones are 
sometimes similar, obscuring bimucronate condition. Hydrotheca length : breadth ratio 
usually 3^t but Millard (1966) gave 1|-3|, once 1$ (BMNH 1935.7.10.1). Angle of slope of 
diaphragm variable. 


DISPERSIVE STAGE. A medusa. Advanced embryos still in the gonotheca have been widely 
reported as having 4 tentacle buds, as they have in other Clytia species, but the free medusa 
has yet to be described. See also Dispersive Stage under Clytia linearis. 


DISTRIBUTION. A widespread, warm water species recorded in the N Atlantic north to 
Suffolk, SE England. However, C. paulensis has been found at only three English localities 
(present material). Fertile material was reported as 'very common' near St Malo, NW France 
(Philbert, 19356) but all the English material has been infertile. 


Table 3 Clytia paulensis Measurements in //m. 

NW France NW France South Africa 

(1937.7.10.1) (Philbert, 19356) (Millard, 1966, 


S Indian Ocean 
(holotype; Vanhoften, 






Breadth (max) 




Pedicel length 



Gonotheca (?rf = 9) 

Breadth (max) 
Aperture diameter 
Pedicel length 


Other European records are 'the Mediterranean' (Picard, 19586), Naples (Riedl, 1959) and 
the Glenan Isles, NW France (Fey, 1969); but not the Roscoff area (Teissier, 1965; L. 
Cabioch, pers. comm.). 

HABITAT. Lower shore (present material) to 200m (Stechow, 19230; Riedl, 1959; 
Mediterranean); 0-138 m, once 384 m (Millard, 1975, 1977; S Africa). Tolerant of reduced 
salinity at least as low as 17%o (Calder, 1976). Usually recorded epizoic on hydroids and 
other inert animal substrates. The following have been reported: Laomedea sp., Clytia 
hemisphaerica (as C. gracilis), Dynamena sp., Halecium beanii Johnston, 1838, Nemertesia 
antennina (Linnaeus, 1758), Pennaria disticha sensu Brinckmann-Voss, 1970, Sertu- 
larella sp. and spines of an echinoid, Cidaris sp. (as Dorocidaris sp.) (all by Stechow, 
19230); Sertularia cupressina Linnaeus, 1758, and Halecium beanii (by Philbert, 19356); 
Hydrallmania falcata (Linnaeus, 1758), Obelia dichotoma and a bryozoan, Crisia sp. 
(present material); and Nemertesia sp., Tubularia sp. and Scrupocellaria scruposa (Bryozoa) 
in Suffolk and Essex, SE England (by R. G. Hughes, pers. comm.). 

REMARKS. The four tentacle buds of the pre-release medusa and the close similarity to C. 
hemisphaerica indicate that paulensis is correctly referred to Clytia. C. paulensis was placed 
in Obelia by Naumov & Stepanyants (1972) because the material they saw was 'strongly 
branched' but this seems slim reason. Stepanyants (1979) later referred C. paulensis to 
'Obelia bicuspidatd* Clarke, 1875, a conclusion with which I cannot agree (see notes under 
O. bidentata, p. 1 1 7). 

The earlier Torbay material listed above, collected in 1973, was the first to be recorded 
from the British Isles. E. T. Browne's Plymouth material had been collected in 1899 but was 
overlooked. Browne first identified it as Campanularia raridentata Alder, here referred to 
Clytia hemisphaerica; but in 1927 added a note doubting his determination (E. T. Browne 
ms notebooks 13 : 148-149; Zoology Library, BMNH). The material came to the BMNH as 
part of the E. T. Browne bequest and was re-identified by the then curator as 'Clytia gracilis 
Sars'. The hydrothecal rims, with their rounded bimucronate cusps, clearly differ from those 
of C. gracilis s. str. auct. and the specimen resembles C. paulensis in all respects. Although 
Browne regarded the material as something unusual in 1899 C. paulensis was not described 
until 1910, and he may well have been the first in the World to collect this species. 

The type material of Clytia ulvae Stechow, 19190, from Marseille, has a bimucronate 
hydrothecal rim and is undoubtedly C. paulensis. Nevertheless, Stechow was among the first 
to collect C. paulensis from Europe and to recognize it as different from C. hemisphaerica. 

Clytia paulensis is perhaps difficult to identify. It differs from C. hemisphaerica in its 


bimucronate hydrothecal rim, its slender hydrotheca and pedicel and, so far as is known, its 
consistently smooth gonotheca. Both species are widespread in the World. But in European 
waters C. paulensis has yet to be found north of Suffolk, SE England, whereas C. 
hemisphaerica ranges much further north. Other west European Campanulariidae having 
bimucronate hydrothecal rims are Obelia bidentata and Laomedea neglecta which produce 
taller colonies and branch differently. 

Clytia incertae sedis 

Two Clytia medusae, with 5 and 6 radial canals and diameters of 8 mm and 1 3 mm respec- 
tively, were taken in the southern Adriatic Trough on 9 August, 1969. They were referred to 
Phialidium pentata Mayer, 1900, by Schmidt & Benovic (1977), making the first NE 
Atlantic record of that species. Kramp (1961) had referred "P. pentata' to Phialidium 
folleatum McCrady, 1857 (= Clytia folleata) but there remains a possibility that both are 
abnormal variants of C. hemisphaerica. Indeed, Schmidt & Benovic considered: 'It might be 
possible that all [recorded] specimens [of pentata, folleata and also P. gardineri Browne, 
1904] are abnormal forms of C. hemisphaerica'. I concur with their conclusion that more 
material is needed to resolve these problems. 

Leloup (1940 : 21, as Laomedea) recorded Campanularia kincaidi Nutting, 1899, from 
the Azores, at 1 187 m depth, without description or comment. The record was repeated by 
Rees & White (1966:277, as Obelia). The only previous record from the Atlantic 
was of two colonies from the Caribbean, also by Leloup (1935 : 20). In the absence of more 
definite indication it seems best to omit the species from the present survey. The nominal 
species was provisionally referred to Clytia by Cornelius (19750 : 280). 

Subfamily OBELIINAE Haeckel, 1879 

Obelidae Haeckel, 1879 : 163 (part). 

Obelinae: Mayer, 1910 : 231 (part); Russell, 1953 : 296. 

NOMENCLATURE. The root of the subfamily name is Obelia, and the spelling Obelinae is 

DIAGNOSIS. Campanulariidae with erect hydrocaulus and true hydrothecal diaphragm; no 
sub-hydrothecal spherule; stolon not anastomosing; medusa liberated but reduced (Obelia), 
or vestigial and retained (the rest). 

TYPE GENUS. Obelia Peron & Lesueur, 1 8 100 (by present designation). 

SCOPE. The genera Gonothyraea Allman, 18640, Hartlaubella Poche, 1914, Laomedea 
Lamouroux, 1812 and Obelia Peron & Lesueur, 18100. 

REMARKS. The subfamily name is the oldest available. The Obeliinae was recognized also by 
Mayer (1910) and Russell (1953), who like Haeckel based their classifications on the medusa 
stage alone. All the included genera occur in the eastern North Atlantic and are defined 

Genus GONOTHYRAEA Allman, 18640 

Gonothyraea Allman, 1864a : 374. 

Gonothyrea auct. (laspus pro Gonothyraea). 

Campanularia, Laomedea and Obelia part, auct. (see Remarks). 

TYPE SPECIES. Laomedea loveni Allman, 1 8590; designated by Millard (1975). The originally 
included species were L. loveni, Campanularia geniculata sensu Lister (= G. loveni; see 
Cornelius, 19770 : 47) and L. gracilis Sars. The last named was based on a mixed series 
comprising what was probably Clytia hemisphaerica (hydroid) and G. loveni, but following 
designation of lectotype material (p. 94) it is now subjectively referred solely to C. 



DIAGNOSIS. Campanulariidae forming upright, branched colonies; stem flexuose; hydro- 
thecae tubular, pedicellate, alternate; diaphragm present; no sub-hydrothecal spherule; 
gonophore a gonomedusa. 

REMARKS. I have discussed elsewhere the past confusion between G. loveni and nominal 
species of Campanularia, Laomedea and Obelia (in Cornelius, \911a). This confusion 
probably delayed by some decades the discovery of the alternation of generations in 
medusoid coelenterates; and, it can be claimed, in other animal groups. 

Gonothyraea loveni (Allman, 1 859a) 
(Fig. 15) 

Laomedea gracilis Sars, 1857: pi. 2, fig. 4 only (not pp. 51-54, nor figs 1-3, 5, = Clytia 

hemisphaerica, q.v.). 

Laomedea loveni Allman, 1859a : 138-140. 
Gonothyraea (Laomedea) loveni: Allman, 18646 : 376. 

Gonothyraea hyalina Hincks, 1866 : 297-298; Hincks, 1868 : 184-185, pi. 35, fig. 2. 
Gonothyraea loveni: Hincks, 1868 : 181-183, pi. 25, fig. 2. 
Obelia loveni: Naumov, 1960 : 264-265, fig. 152 (syn. G. hyalina Hincks); Naumov, 1969 : 285-287, 

fig. 1 52 (syn. G. hyalina Hincks). 

NOMENCLATURE. Bedot (1912 : 294; 1916: 107) listed but two uses of the combination 
Obelia hyalina, and none of O. loveni, in his synoptic works (1901-1925). Obelia hyalina 
Clarke, 1879, and O. hyaliana Vannucci, 1955, are different nominal species. 


Figs 15-16 Fig. 15 Gonothyraea loveni. (a) tip of colony, Bay of Biscay; 1959.9.17.59. (b) same, 
hydrotheca. (c) 9 gonotheca and gonomedusa, Vadso, E Finmark, Norway, intertidal; 
1912.12.21.184. Scales (a-c) 500 urn. Fig. 16 Hartlaubella gelatinosa. (a) part of 9 colony. Note 
large embryos. SW England; 1959.9.17.57 (microslide preparation), (b) same, two blastomeres. 
Note conspicious chromosomes (see Remarks), (c) hydrothecal rims. Israel; 1932.8.13.1. (d) 
unopened rf gonotheca, NE England; 1 969. 1 1 .28.2. Scales: (a, d) 500 //m; (b-c) 50 /^m. 


TYPE LOCALITY AND MATERIAL. Firth of Forth, Scotland (Allman, \S59a : 137); material not 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Gonothymea hyalina Hincks, 1866, syntypes, 
Shetland Is, Scotland, several colonies in spirit, coll. J. G. Jeffreys, on 'Tubularia, 
Halecium,etc.\ex Hincks colln; 1899.5.1.157. 

OTHER MATERIAL EXAMINED. BMNH collection, about 80 specimens. 

DESCRIPTION. Colony erect, stem monosiphonic, delicate, variably flexuose, internodes 
usually slightly curved, irregularly branched, up to c. 100mm. Annuli above origins of 
branches. Hydrothecal pedicels usually annulated throughout but smooth central portion 
frequent; usually tapering distally to roughly half proximal diameter. Hydrotheca 
cylindrical, campanulate, length 1^-3 times maximum breadth; rim delicate, slightly out- 
turned, castellate, raised portions often notched (Fig. 15); often longitudinal folds in 
hydrotheca. Hydranth with 20-25 tentacles; branched tentacles recorded but rare 
(Hamond, 1957); hypostome spherical. Gonotheca (rf narrower than 9, otherwise similar; 
Miller, 1973) cylindrical, truncated above, tapering below. Conspicuous sporosacs or ova 
develop internally and later extrude up to four together as tentaculate, sub-spherical reduced 
medusae termed gonomedusae (formerly called meconidia; see Dispersive stage). 
Variation. Internode length and curvature, length of hydrotheca and the degree of tanning 
and translucency of the perisarc are all variable. 

DISPERSIVE STAGE. A planula larva. The reduced medusa is retained until after the planulae 
have escaped. It does not swim but simply drops off. Ellis (17560, b, c, 1767, but not 1755; 
see Cornelius, \911a) recorded that the 'released' gonomedusae exhibited strong tentacle 
movements and adopted a 'worm-like' shape, but his observations seem not to have been 

The gonomedusae were interpreted as highly developed sporosacs by Allman (1 859a) who 
coined for them the long standing term meconidia. But Goette (1907) and more recently 
Miller (1973) have shown that they are actually reduced medusae. Miller introduced the 
preferable term gonomedusae. Wulfert (1902) provided a summary of early reproductive 
studies on this species. 

REPRODUCTIVE SEASON. Published records suggest that in the English Channel and southern 
North Sea the species breeds almost throughout the year (Mar-Sep, Hamond, 1957; Jan-Apr 
& Sep-Nov, Marine Biological Association, 1957; May-Aug & Oct-Nov, Teissier, 1965); 
but an intertidal population which I studied in 1974 in Sussex, SE England, bore 
gonomedusae only during the first two weeks of April. Some of the published records might 
refer to colonies with developing gonothecae, or with empty ones. 

Fertile gonothecae were reported at Naples from January to May and in September by Lo 
Bianco (1909). 

DISTRIBUTION. Widespread in suitable habitats and often common. Occurs throughout 
western Europe north to W Greenland (but not E), Iceland, Faeroes, Spitzbergen, Barents 
Sea, Murman coast and White Sea (Mathiesen, 1928; Kramp, 1929, 1938; Calder, 1970). In 
the Baltic G. loveni is said to penetrate as far as Helsinki in the Gulf of Finland and the Aland 
Isles in the Gulf of Bothnia (Linko, 1911; Stechow, 1927; Naumov, 1969). It has been 
recorded from the Mediterranean Sea (Picard, 19586; Riedl, 1959) and Black Sea (Naumov, 
1969); and on the Atlantic coast from W France and Morocco (Billard, 1927; Patriti, 1970). 

Millard (1975) recorded the species in South Africa only from Cape Town docks and 
considered this and other Southern Hemisphere records to result from transport by ships. 
Hence G. loveni might be found further south than Morocco. Rees & White (1966) listed a 
dubious old Azores record from the unusual depth of 845 m. 

HABITAT. Intertidal, usually in pools, and offshore. Naumov (1960, 1969) gave a normal 
depth range of 0-30 m, with an extreme lower limit of 200 m; and Mathiesen (1928) 


similarly gave 6-200 m. The BMNH material falls within these limits. The record at 845 m 
off the Azores listed by Rees & White (1966) is much deeper, and needs confirmation. G. 
loveni is tolerant of reduced salinity at least to 12%o (Calder, 1976). It has been recorded on a 
variety of animal, plant and inorganic substrates (Hincks, 1868; Hamond, 1957; Marine 
Biological Association, 1957; Teissier, 1965), and there is no regular association. 

REMARKS. Hummelinck (1936), Naumov (1960, 1969) and others listed by Calder (1970) 
regarded Gonothyraea loveni and G. hyalina as conspecific and I concur. 

The distinctive method of reproduction has been described in part by many authors from 
Ellis onwards. Nevertheless G. loveni was widely confused with Obelia spp. and with 
Laomedea Jlexuosa until Wright (1858, 1859) realized it was distinct (Cornelius, 19770; 
Cornelius & Garfath, 1980). But Wright did not provide a name. The species was soon 
named by Allman (18590) who took Joshua Alder's suggestion (in litt.) that the species was 
valid. Alder was perhaps unaware of Wright's publications. From the literature it would 
appear that Wright and Alder worked independently; but since both communicated freely 
with Hincks (e.g. 1 868 : preface) they could have been in touch directly. It may be difficult to 
find out who among these British workers really was first to recognize G. loveni. 

Most of the life cycle is well understood and has been redescribed in detail by Miller (1973; 
see also Bergh, 1879, Wulfert, 1902, and Goette, 1907). Aspects of stolon growth were 
described by Wyttenbach, Crowell & Suddith (1973). 

Laomedea gracilis Sars, 1850, 1857, was based on a mixed type series and this has caused 
confusion. The material illustrated in 1857 was partly G. loveni (see synonmy) but 
predominantly Clytia hemisphaerica (see Stechow, 19230 : 111). The material resembling C. 
hemisphaerica in this series is here designated lectotype. Hence the species name loveni 
Allman, 18590, remains available. In any case, the name gracilis is preoccupied. This and 
other problems relating to "L. gracilis'' are considered under C. hemisphaerica, where a new 
name is introduced in place of L. gracilis (p. 78). 

Obelia hyalina Clarke, 1879, has been referred to Obelia dichotoma (by Cornelius, 
19750 : 266) and is not G. hyalina Hincks. Hence Billard's (19310) record of 'Obelia hyalina 
Clarke' from Mauritania refers to O. dichotoma and not G. loveni. This homonymy is 
discussed further under O. dichotoma (p. 1 19). 

The affinities of G. loveni are discussed above (pp. 47-49). 

Genus HARTLAUBELLA Poche, 1914 

Sertularia, Campanularia, Obelia and Laomedea auct., part. 
Obelaria Haeckel, 1879 : 172 (part). 
Obelaria Hartlaub, 1897 : 488 (homonym). 
Hartlaubella Poche, 1914 : 76. 

TYPE SPECIES. Sertularia gelatinosa Pallas, 1 766; by monotypy. 

DIAGNOSIS. Erect, colonial Campanulariidae with polysiphonic stems and second order 
branching; hydrotheca pedicellate, with diaphragm and castellated rim, without spehrule; no 
medusa stage (see Remarks under H. gelatinosa). 

REMARKS. The genus name Obelaria Haeckel, 1879, was proposed as a nom. gen. nov. for 
the hydroid stage of Obelia, of which name it is a junior synonym (Cornelius, 19750 : 254). 
Haeckel included Sertularia gelatinosa Pallas, 1 766, in its scope. Confusion was caused later 
when Hartlaub (1897) independently coined the generic name Obelaria to apply to a gen. 
nov. comprising Sertularia gelatinosa alone. Although Hartlaub's name is a homonym of 
Haeckel's and cannot be used, Hartlaub's generic concept is accepted here. The next avail- 
able name for the genus is Hartlaubella Poche, 1914, of which gelatinosa has always been 
the only member. 


Hartlaubella gelatinosa (Pallas, 1 766) 
(Fig. 16) 

Sertularia gelatinosa Pallas, 1 766 : 1 16-1 1 7. 

Campanulariaflemingii Deshayes & Edwards, in Lamarck, 1836 : 133-134. 

Laomedea gelatinosa: Couch, 1884 : 4-5, 39-40; Hincks, 1852 : 85-86; Da Cunha, 1944 : 65-66; 

Vervoort, \946a : 300-303, fig. 133 (syn. Campanulariaflemingii Deshayes & Edwards). 
Obelia gelatinosa: Hincks, 1868 : 151-154, pi. 26, fig. 1; Naumov, 1960 : 268-269, figs 157-158; 

Naumov, 1969 : 290-29 1 , figs 1 57-1 58. 
Obelaria gelatinosa: Haeckel, 1879 : 172, 173, 176; Hartlaub, 1897 : 488-495 (non Haeckel); Nutting, 

1915: 88-90, pi. 24, figs 1-5. 
Hartlaubella gelatinosa: Poche, 1914:76; Stechow, 1925:522; Stechow, 1927:309; Teissier, 

1965: 17;Calder, 1970: 1 543; Cornelius &Garfath, 1980 : 283. 
Campanularia gelatinosa: Ralph, 1957 : 820, fig. Ib-f. 

TYPE LOCALITY AND MATERIAL. Belgian coast (Pallas, 1766; Ralph, 1957); specimen not 

MATERIAL EXAMINED. BMNH collection, about 50 specimens. 

DESCRIPTION. Colony elongate; small colonies loosely conical, large colonies bushy, up to c. 
200 mm; main stems straight or branched, polysiphonic. Final branches monosiphonic, 
arranged irregularly all round stem, dichotomous, flexuose; internodes usually curved but 
sometimes straight, length varied, 5-9 rings basally. Hydrothecae on ringed, slightly tapering 
pedicels, long-campanulate to cylindrical; rim often abraded even but initially castellate 
with notch of varied depth in centre of each blunt cusp; embayments rounded; diaphragm 
transverse. Hydranth with 22-28 tentacles, hypostome spherical. Propagation by stolons in 
spring. Gonotheca ^ = 9, axillary, inverted-conical, sides smooth to sinuous; aperture wide, 
distal, on short collar; ova (4^6) and embryos larger than usual in family; nuclei of 
blastomeres large, with unusually conspicuous chromosomes. 

Variation.lntQrnode length and curvature, angle of flexure of stem and length : breadth ratio 
of hydrotheca are all variable. The hydrothecal rim often abrades smooth. 

DISPERSIVE STAGE. Planulae, developing within the gonotheca. Some authors have 
mistakenly reported a medusa stage. 

REPRODUCTIVE SEASON. May-August in NW France (Teissier, 1965); July, 1934, 
Northumbria (H. O. Bull, in Evans, 1978). 

DISTRIBUTION. Recorded from southern Scotland, Oslo Fjord, Danish waters and part of the 
Baltic south to the Mediterranean and Black Seas. The species is common in Dutch, Belgian, 
Irish, Welsh, English and N & W French waters (Vervoort, 19460; Leloup, 1952; BMNH 
collection; Billard, 1927; Teissier, 1965). 

Trustworthy Scottish records are few and there seems only one this century, although the 
species is still common in NW England: Tay Estuary (Fleming, 1820; Alexander, 1932); 
Shetlands, Berwick Bay and Solway Firth (Johnston, 1847); ?Dundee (BMNH 
1851.7.25.227, specimen not located). Hincks (1868) repeated several of these records and 
Norman (1869) gave another Shetlands locality. Recent English records are numerous, 
northerly ones including Northumberland (Alexander, 1932) and Morecambe Bay (J. Clare, 
pers. comm. & BMNH 1970.8.28.6-10). 

The species was recorded from 13 localities in Oslo Fjord by Christiansen (1972) but his 
statement that the species occurs north to Finmark is questionable, and Linko (1911) and 
Broch (1918) listed no records so far north. Neither also did Kramp (1929, 1938), who 
likewise considered the species absent from Iceland and the Faeroes. Naumov (1969) gave an 
Iceland record without further detail; but with greater precision stated the northern limit in 
the Baltic to be Tort Liepaja = Libava', Latvia. Kramp (1935) listed several Danish records 
north to Frederikshavn, NE Denmark, and repeated Stechow's (1927) record from Trave- 
miinde, near the southern limit of the Baltic Sea. Rasmussen (1973) reported the species 
from Siaelland Island, southern Denmark. 


Mediterranean records are few. Naumov (1969) gave 'Black Sea, Mediterranean and 
Gibraltar'. Although Picard (19586:197) expressly excluded the species from the 
Mediterranean list Rossi (1950 : 205) had earlier recorded it from NW Italy, the Adriatic and 
Strait of Gibraltar. Linko (1911) listed Black Sea material. 

The species is known from Portugal (Nobre, 1931; Da Cunha, 1944) but has yet to be 
recorded so far south as Morocco (Patriti, 1 970). 

HABITAT. Intertidal, particularly in pools and run-off from saltings, and shallow depths off- 
shore. Often in estuaries and tolerant of reduced salinity at least to 6 - 2%o(Vervoort, 1946a). 
Also tolerant of silt. 

The lower depth limit is probably c. 15 m. A record from 30 m off the Scilly Isles was 
based on Obelia bidentata material (BMNH 1969.9.9.6; mentioned, Robins, 1969). Couch 
(1844) recorded material from beneath intertidal stones and on algae, but that under stones 
might have been Laomedea neglecta (cf. p. 107). 

REMARKS. The nomenclatural history of this species is involved. The name Sertularia 
gelatinosa was first applied by Pallas (1 766) to the 'Corallina confervoides, gelatinosa alba, 
geniculis crassiusculis, pellucidis' of Ray (1724 : 34, para. 7). However, Ellis (1755 : 20, 
p. 1 1 , figs B, b) assigned bryozoan material to Ray's species, providing clear illustrations. 
Linnaeus (1758 : 812) included the descriptions of both Ray and Ellis in a single species to 
which he gave the new name Sertularia spinosa. This name is currently applied to a 
bryozoan species in the combination Vesicularia spinosa (Linnaeus, 1758), for example by 
Prenant & Bobin (1956). The bryozoan name Sertularia sericea Pallas, 1776 : 1 14, was a 
nom. nov. for S. spinsoa and is its junior objective synonym. Pallas elsewhere 
(1 766 : 1 16-1 1 7) adequately described gelatinosa. 

Fleming (1820) noted that Ray and Ellis each described a different species but referred 
Ray's description to Obelia geniculata, not to the present species. However, Pallas' account 
is clear and he contrasted gelatinosa with both O. geniculata and O. dichotoma. Also he 
noted the polysiphonic stem and cusped hydrothecal rim both unusual in Obelia. Even so, 
it could be argued that Pallas had material of Obelia bidentata before him since that species 
superficially resembles H. gelatinosa. There is some evidence that O. bidentata did not then 
occur in European waters. Even if it did, Pallas might have overlooked the fine cusps on the 
hydrothecal rim which are a main distinction. But despite these small doubts it seems highly 
likely that Pallas' description indeed refers to the present species. Hincks (1868: 152) 
himself commented that Pallas' description was 'admirable, and is the only one we possess 
which is not positively incorrect'; so the case is strong. Hincks summarized some of the 
additional taxonomic confusion surrounding the species between 1 820 and 1 868. 

Campanularia flemingii Deshayes & Edwards, in Lamarck, 1836, was based on material 
of the present species described by Fleming (1820) from Scotland although Deshayes & 
Edwards gave the type locality as coasts of England! Fleming had referred his material to 
gelatinosa but noted that it disagreed with Pallas' description in having even hydrothecal 
rims. Fleming thought Pallas might have mistaken tentacle tips for castellations on the 
rim, and assumed that gelatinosa always had an even rim. Deshayes & Edwards thought 
Pallas too careful to make this mistake and concluded that two species were involved, one 
with castellations and one without. Probably Fleming's material simply had hydrothecae 
in which the rims were worn smooth! Johnston (1838, 1847) realized the confusion and 
referred flemingii back to gelatinosa. He was followed by Bedot (1905) and Vervoort 
(1946a), and I concur. Gray (1848), however, gave C. flemingii specific rank, but did not cite 
material. Gray seems usually to have relied heavily on Johnston's work but on this occasion 
clearly did not. Possibly Edwards, who sometimes worked on the British Museum collec- 
tions, persuaded him to accept the species. 

Thaumantias leucostyla Will, 1844 : 73, pi. 2, figs 16-17, based on an Obelia medusa, was 
referred to the present species by Bedot (1912 : 328) without comment; but as gelatinosa has 
no medusa stage this must be wrong (see next paragraph). It would be difficult to identify the 
medusa beyond Obelia sp. from Will's description. 


Several authors, including Hincks (1852, 1868), have stated that H. gelatinosa releases a 
medusa, which it does not (Cornelius, \915a : 279). It could be that Hincks and the others 
saw medusa release from Obelia bidentata. But the first European records of that species date 
from the early 1900s; and it seems improbable that Hincks would have made such a mistake 
when identifying a hydroid. His comment that the branches of gelatinosa 'droop slightly' fits 
bidentata, but this is not conclusive. Another possible explanation of his apparent mistake is 
that he took the unusually large ova of//, gelatinosa for developing medusae. 

Couch (1844) had much earlier given a correct (and posthumous) description of the life 
cycle, reporting planula development and release, and early development of the young 
colony. In addition he noted that the planulae were propelled by cilia. But his contempor- 
aries were still muddled, and had Couch lived a little longer he might have corrected some of 
the ensuing confusion. Van Beneden (1843, 1844) reported medusa release in H. gelatinosa 
but his material was actually Obelia dichotoma (see Cornelius, 19750; and 19770 for other 
references). Hincks (1852) also attributed a medusa to gelatinosa. Van Beneden's error of 
identification was later appreciated by Hincks, who referred Van Beneden's material to 
a synonym of O. dichotoma, namely O. longissima (Pallas, 1766). Similarly, the much later 
report by Godeaux (1941) that gelatinosa had a medusa was also based on O. dichotoma 
material. It might be relevant that Godeaux worked at the Van Beneden Institute! Then 
Leloup (1947), paralleling Hincks, referred Godeaux' material to O. longissima. 

A convenient distinction between O. bidentata and H. gelatinosa is that in side view the 
branches of bidentata show a graceful sigmoid curvature lacking in gelatinosa. 

The large chromosomes illustrated here recall in shape and size those of Obelia medusae, 
shown by Faulkner ( 1 929). 

Genus LAOMEDEA Lamouroux, 1812 

Laomedea Lamouroux, 1812 : 184. 

Campanularia Lamarck, 1816 : 112 (part). 

Lomedea Pickering, in Dana, 1846 : 689 (lapsus pro Laomedea, see p. 78). 

Campalaria Hartlaub, 1897 : 449. 

Eucampanularia Broch, 1910: 1 84 (part; see p. 52). 

Eulaomedea Broch, 1910 : 189; Millard, 1975 : 223. 

Cmpanularia Mulder & Trebilcock, 1914: 11 (part; lapsus pro Campanularia). 

Laomedea (Paralaomedea) Hummelinck, 1936 : 57; Vervoort, 19460 : 285. 

Eulaomeda Rees & Thursfield, 1965 : 102 (lapsus pro Eulaomedea). 

TYPE SPECIES. Laomedea flexuosa Alder, 1857 (proposed designation by Cornelius, 1981). 
Broch (1905 : 10) proposed 'Laomedea loveni Allman, 18590' as type species but loveni was 
not among the species originally included in the genus and is not eligible. 

DIAGNOSIS. Colonial Campanulariidae with: polyp generation forming upright colonies; 
stolon branching but not anastomosing; hydrotheca pedicellate, lacking spherule; true 
diaphragm present; annular perisarc thickening inside base of hydrotheca; gonotheca 
stolonal or axillary, aperture typically circular, wide; gonophores sessile, interpreted as 
vestigial medusae in many species. 

REMARKS. I have previously pointed out (Cornelius, 19750) that Laomedea Lamouroux, 
1812, is a junior synonym of Obelia Peron & Lesueur, 18100. But the name Laomedea is so 
well known that I have submitted a case to the International Commission on Zoological 
Nomenclature recommending that it be conserved by application of the Plenary Powers. I 
have proposed that Laomedea flexuosa Alder, 1857, be designated type species although it 
was not originally included. The alternative, of applying the Rules of Nomenclature, would 
result in the virtually unused name Campalaria Hartlaub, 1897, being employed for the 
present genus. Further details of the case, and another concerning the genus name 
Campanularia (p. 5 1 ), have been presented elsewhere (Cornelius, 1981). 
The subgenus name Eulaomedea Broch, 1910, type species Laomedea flexuosa Alder, 


1857, by monotypy, is a junior synonym. Stechow (1923a : 95) referred Eulaomedea to 
Laomedea Lamouroux, 1812, and apart from subgeneric use by Splettstosser (1924), 
Hummelinck (1936) and Vervoort (1959) there seems to have been no further appearance of 
Eulaomedea in the literature until Rees & Thursfield (1965). These authors upgraded 
Eulaomedea to genus status, including within its scope "E. angulata\ 'E. flexuosa" 1 and '. 
calceolifera\ Finally Millard (1975) employed Eulaomedea to embrace flexuosa and 
calceolifera alone. Thus the name has not been widely used and Broch (e.g. 1918) himself 
came to drop it, without comment. For further details see Cornelius (1981). 

The subgenus Paralaomedea was apparently introduced by Hummelinck (1936). The type 
species is Laomedea neglecta (Alder, 18566; by monotypy). Vervoort (19460) followed 
Hummelinck's usage. But the unusual acrocyst of neglecta was shown by Splettstosser 
(1924) to be medusoid in origin, so there seems inadequate reason for subgeneric separation. 

Laomedea angulata Hincks, 18616 
(Fig. 17) 

Laomedea angulata Hincks, 18616 : 261, pi. 8; Hummelinck, 1936 : 51-52, fig. 5; Picard, 19586 : 191 
(syn. L. sphaeroidea Stechow); (non Da Cunha, 1944 : 63, fig. 36; = Laomedea calceolifera). 

Campanularia angulata: Hincks, 1868: 170-171, pi. 34, fig. 1, woodcut 14 (p. 136); Fraipont, 
1880 : 433-466, pis 32-34; Billard, 19046 : 46, 47, 53, 55, 57, 65, 67, 72-82, 97, 144, 173, pi. 3, figs 
1-7, pi. 5, figs 1-2, 7, 10; Faure, 1965 : 419-426, figs Ib, d, 2a, b, 3a, b. 

Laomedea sphaeroidea Stechow, 1932 : 85-86. 

Eulaomedea angulata: Rees & Thursfield, 1965 : 101-102. 

TYPE LOCALITY AND MATERIAL. Hincks (18616) based the original description on specimens 
from South Devon and the Isle of Man. Some of this material is preserved as follows: (i) 
Hancock Museum, Newcastle upon Tyne, infertile colony on single blade of Zostera L. (eel 
grass), in spirit, labelled 'Laomedea angulata, Ramsey, Isle of Man. Revd T. Hincks' and, on 
a second label, 'Campanularia angulata Hincks' (mentioned, Cornelius & Garfath, 1980); 
(ii) BMNH, several colonies on blades of Zostera, in spirit, in two tubes. One contains a 
single, wide blade of Zostera supporting an infertile colony of C. angulata; the other tube 
five Zostera blades, all much narrower than that in the first, each supporting one or more 
fertile colonies of C. angulata. The two tubes, in one jar, jointly bear the registered number 
1899.5.1.149. In the bottom of the jar, detached from the tubes, were two faded labels written 
by Hincks: 'Campanularia angulata Hincks, Isle of Man', and ' Campanularia angulata 
Hincks, Britain'. There seems no indication which label originally belonged to which tube. 
Although the narrow leaved Zostera resembles that in the Hancock Museum specimen, 
labelled Isle of Man, there is no further indication that the BMNH thin leaved specimen 
came from there too. 

Probably all this material was identified by Hincks at some time, but whether he saw it 
before or after the date of publication of the first description is not clear. Hence, it is not 
possible to decide whether the specimens should be treated as syntypes or neotypes. 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Laomedea sphaeroidea Stechow, 1932, fertile 
colonies taken from Zostera, Sete, S France, microslide preparations; Munich Zoological 

OTHER MATERIAL EXAMINED. BMNH collection, c. 30 specimens from Britain and the 
Channel Islands and 13 microslide preparations from the Lagoon of Venice. Amsterdam 
Zoological Museum and Leiden Natural History Museum, Netherlands, c. 20 specimens. 

DESCRIPTION. Mature colony comprising several erect, monosiphonic stems inserted at 
approximately regular intervals on an almost straight, little branched or unbranched, smooth 
hydrorhiza; recorded reliably only on eel grasses; stolons usually (Fraipont, 1880; Billard, 
19046) growing parallel with the leaf margins. Stems flexuose, internodes markedly straight 
(rarely, faintly curved), angle between them 90-120, length : breadth ratio variable; \-c. 10 
annuli basally in BMNH material. Late in season stems terminate in long, often curling 



Fig. 17 Laomedea angulata. (a) part of colony, including hydrocaulus and stolonal 9 gonotheca 
with embryos at different stages of development, Jersey; 1959.9.17.11. (b-c) cf gonothecae, 
Plymouth, SW England; 1959.9.17.3, 12. (d) vegetative terminal stolon; as (c). Scale (a-d) 
500 /on. 

tendrils approximately same width as internodes except at often recurved tip region which is 
wider. Tendrils probably represent modified hydranths: those ofObelia dichotoma differ (see 
Remarks). Hydrothecae campanulate, delicate, usually slightly flared distally, length c. \{ 
times maximum breadth, thin walled and not thickened; diaphragm usually transverse but 
occasionally slightly oblique; pedicels 3-1 5 ringed, slightly tapering distally, sometimes with 
smooth central portion (e.g. BMNH 1959.9.17.11). Hydranth with 24-30 long tentacles 
alternately elevated and depressed; hypostome large, spherical when dead but (Fraipont, 
1880) when alive constantly changing in shape; tissues colourless. Gonothecae apparently 
always borne on stolon. 9 elongate-ovate, aperture distal, wide, on slightly demarcated neck, 
usually with sub-terminal internal strengthening ring; on short, 3-6 ringed pedicel; contain- 
ing several eggs (see Remarks); planulae brooded internally. <5 similar, aperture narrower; 


containing several cf gonophores; sometimes said to be more tapered distally than 9, but this 
not evident in BMNH series. No medusa stage. 

Variation. Apparently minor. Broch (1933) regarded L. calceolifera a variety of the present 
species, proposing a forma typica for L. angulata s. str., but his opinion is no longer followed. 

DISPERSIVE STAGE. Planulae, which develop within the gonotheca (e.g. BMNH 
1 959.9. 17.11). Also vegetative tendrils which break away to form new colonies. 

REPRODUCTIVE SEASON. Sexual reproduction June-August in NW France (Teissier, 1965), 
but fertile material recorded early as April in Isle of Man (Bruce et al., 1963, material not 
examined). Vegetative reproduction involving tendrils typically August to November in NW 
France (Teissier, 1965). 

DISTRIBUTION. A southerly species which probably occurs no further north than the British 
Isles. Unchecked published records and data with BMNH specimens indicate the following 
localities in NW Europe: SW Scotland (Rankin, 1 90 1 ; Ritchie, 1911; Chumley , 1 9 1 8); N & S 
Ireland (Hincks, 1868); Isle of Man (part of type series; also Bruce et al., 1963); Scilly Isles, 
1967 (Robins, 1969); S Devon, 1898 & 1906 (type series; also Marine Biological Associ- 
ation, 1957; a 1936 record on a barnacle seems unlikely); Dorset, BMNH; Roscoff(Fraipont, 
1880; Faure, 1965; Teissier, 1965); Channel Islands (Vervoort, 1949; Leiden NHM & 
BMNH); Netherlands (Vervoort, 1946#; Leiden NHM). There seems only a single, 
unchecked record from the east coast of Britain, from St Andrews Bay (Crawford, 1895, 
repeated in Laverack & Blackler, 1974); and the species has not often been recorded from the 
many Zostera beds of southern England (see Addendum). 

Published records from Spain southwards include the following: Santander, N Spain, 
(Rioja y Martin, 1906); NW Italy (Rossi, 1950; also Naples, Riedl, 1959, and lagoon of 
Venice, BMNH material); 'Mediterranean' (Picard, 19586). 

At Castiglione, Algeria, Picard (1955) found no less than 1 8 species of hydroids growing on 
the eel grass Posidonia, but did not report L. angulata. A single colony of the species was 
erroneously reported from Portugal by Da Cunha ( 1 944), his illustration showing a mature 9 
gonotheca of L. calceolifera. The record from the Falkland Islands by James Ritchie, 
repeated by Rees & Thursfield (1965), was based on Obelia dichotoma material. 

The northernmost material I have seen came from the Isle of Man (type series; also Bruce 
et al., 1963). More northerly material was reported from the Clyde Sea by both Rankin 
( 1 90 1 ) and Ritchie ( 1 9 1 1 ), the latter repeating some of Rankin's records and giving some new 
ones. However, the depth range given by the two authors, 30-1 30 m, exceeds the depths from 
which the species has otherwise been recorded. There are records of one of the substrate 
plants, Zostera, from the Clyde Sea area so it is conceivable Rankin and Ritchie at their 
deeper localities dredged up loose plants which had sunk; but dead Zostera often floats! Den 
Hartog (1970) gave 7 m as the deepest British record of Zostera marina L. (in the Scilly Isles) 
but cited Danish and Mediterranean records down to 1 1 m and one from the Pacific coast of 
the U.S.A. at 30 m, adding credibility to at least some of the Clyde Sea records. He gave the 
NE Atlantic distribution of Z. marina as Algeria (Castiglione), S France (rare) and N 
Mediterranean north to the arctic coast of the U.S.S.R., so it can be assumed that the 
northern limit of L. angulata is not determined by availability of eel grass. 

However, the close association of this species with the eel grasses certainly affects its 
distribution. Thus Zostera almost died out in England in the 1930s (Tutin, 1942), and there 
is a virtual lack of L. angulata records since the 1900s. See also Addendum. 

L. angulata was not recorded in several surveys of North Sea coasts, as follows: 
Northumberland and Durham (Robson, 1914), Norfolk (Hamond, 1957; Hamond & 
Williams, 1977), Belguim (Leloup, 1952), Denmark (Kramp, 1935; Rasmussen 1973) and W 
Sweden (Rees & Rowe, 1969). The record from Danish waters by Vervoort (19460) probably 
referred to Broch's (1928) record as Campanularia conferta and should be rejected (W. 
Vervoort, pers. comm.). The species has still to be recorded from Denmark (K. W. 
Petersen, pers. comm.); but a record from the Great Belt, Kattegat, by Winther (1879, 


repeated in Stechow, 1927) if confirmed would be the most northerly of the species. A record 
dated 1960 from 20 m depth off SW Wales, quoted by Crothers (1966), seems dubious since 
both depth and area would be unusual. However, Zostera does occur there (K. Hiscock, pers. 

HABITAT. Intertidal and shallow sublittoral, the deepest reliable record being 6-8 m 
(Studland Bay, Dorset, during or before 1 890, coll. R. Kirkpatrick, pres. F. Beckford; BMNH 
1 899.7.22. 1 ). Deeper records, from the Clyde Sea and off SW Wales, cannot be substantiated. 
Faure (1965) reported that L. angulata grew in a zone on the shore at Roscoff above that 
occupied by L. calceolifera; but the numerous records from shallow offshore localities show 
that L. angulata is not confined to the intertidal zone. A record from 64 fathoms (130 m) 
near the Falkland Is repeated by Rees & Thursfield (1965) was based on Obelia dichotoma 

L. angulata has been widely recorded on the eel grasses, Posidonia and Zostera (Hincks, 
186 16; Fraipont, 1880; Philbert, 19356; Hummelinck, 1936; Vervoort, 19460; Rossi, 1950; 
Riedl, 1959; Faure, 1965) and only seldom on other substances (sertularian hydroids by 
Betencourt, 1888; Laminaria by R. Oppenheim, in Vervoort, 1949; Dictyota dichotoma by 
Philbert, 19356; Balanus improvisus by W. J. Rees, in Marine Biological Association, 1957; 
Fucus by several recorders listed in Teissier, 1965). The overwhelming majority of published 
reports, and all the BMNH, Amsterdam Zoological Museum and Leiden Natural History 
Museum material are from Zostera, however, and it is likely that records on other substrates 
are wrong. Possibly some at least refer to L. calceolifera, since the two species were confused 
for several decades. Although Betencourt noted terminal tendrils on his material it might 
have been Obelia dichotoma which also has tendrils and can look remarkably similar. 

Nishihira (1968) made a detailed study of the hydroids epizoic on Zostera in northern 
Japan, but did not report L. angulata which seems (Stechow, 19236) not to occur there. 
Picard (1955) made a similar study in Algeria, where he found L. angulata absent from the 
Posidonia beds of Castiglione. 

The species is tolerant of brackish conditions. Hummelinck (1936) noted a tolerance of 
9-8%o Cl, a figure repeated by Vervoort 19460) and, incorrectly as salinity, by Naumov 
(1960, 1969). The corresponding salinity figure is 17'7%o. Morri (19790) found L. angulata 
in a range of salinities down to 27%oin Tuscany, Italy. 

REMARKS. Athough Hincks (1868, 1871) and several subsequent authors distinguished 
correctly between the present species and L. calceolifera some later authors (e.g. Babic, 
1912; Broch, 1928, 1933; Vervoort, 19460; Naumov, 1960, 1969) regarded them 
conspecific. A summary of this confusion and a taxonomic assessment of the two species was 
provided by Faure (1965). The main differences are as follows, in approximate order from 
most to least useful. The gonothecae differ both in structure and position (hydrorhizal in L. 
angulata, on the stem and axillary in L. calceolifera). The internodes in angulata are usually 
straight, and curve slightly in most calceolifera specimens. Terminal tendrils are often 
present, particularly in autumn, in angulata but are unrecorded in calceolifera (Obelia 
dichotoma sometimes has them also). Eel grasses are probably the only substrate for angulata 
but calceolifera occurs on a variety of inert, solid substrates. Faure reported angulata growth 
at its peak in summer, and that of calceolifera in spring, at the same place. He reported erect 
stems spaced at 2-5 mm intervals in angulata and often clustered in calceolifera; but the 
extent to which this character in angulata is phenotypic, induced by the elongate shape of 
the eel grass leaves, is not known. Finally, Faure reported that colony height was 5-10 mm in 
angulata, 13-1 6 mm in calceolifera, each stem bearing respectively at maturity 4-6 and 
10-15 hydrothecae. Despite this list of differences it remains difficult to identify some 
infertile material, even with the aid of long reference series; and some young colonies are 
probably impossible to identify. 

A straight hydrorhiza occurs in Obelia geniculata also, in which it can be interpreted as a 
genotypic character adapted to keeping neighbouring, parallel-running hydrohizae ad- 
equately spaced along their whole lengths. O. geniculata occurs on broad thalloid algae, and 


spacing is probably an advantage. L. angulata is unusual among thecate hydroids in being 
closely adapted to its substrate in a number of obvious ways, and may have taken to living 
on eel grasses in relatively recent geological time. It seems likely that since eel grasses are 
themselves quite recent, being derived from more conventional terrestrial angiospermes, 
other campanularian and laomedean hydroids were living on algal substrates long before L. 
angulata or its ancestors took to eel grasses. Maybe hydroids of this group, having straight 
hydrorhizae, were pre-adapted to growing on the long narrow leaves of eel grasses. 

However, at least some orientation of stolon growth occurs in L. angulata in a direction 
parallel with the eel grass leaf since diagonal or transverse stolons do not occur. O. geniculata 
stolons, although straight, are not orientated along the lamina and it may be that angulata 
stolon tips have a geotropic response which keeps them growing vertically (upwards or 
downwards) along the leaf, which floats upright when the tide is in. The fact that stolons of 
angulata grow sometimes along the narrow edges of the substrate leaves suggests that the 
planula does not seek a central position; or that once a tendril attaches and forms a new 
colony the hydrorhizal tip cannot locate the centre of the leaf; but there is no experimental 

The terminal tendrils of L. angulata were found to be most common from August to 
November at Roscoff by Faure (1965). Billard (19046) suggested that their function was to 
attach to adjacent eel grass leaves and hence enable the species to colonize fresh plants 
vegetatively. The tendrils of BMNH specimens differ from those of occasional Obelia 
dichotoma colonies in having dilated recurved end regions, the ends of O. dichotoma tendrils 
being uniform in width and approximately straight. Study of the BMNH material suggests 
that tendrils in L. angulata are modified hydranth/hydrotheca complexes. 

The most detailed histological and general biological account of L. angulata is that of 
Fraipont (1 880). However, he stated that the female gonophore contains a single egg; but one 
of his illustrations (pi. 34, fig. 3) showed a female gonotheca containing several planulae, as 
illustrated also here, and more than one egg seems normal. 

The nominal species Laomedea sphaeroidea Stechow, 1932, was based on material from 
near Sete (Cette), S France, which Stechow (\9\9a) had earlier referred to L. angulata. The 
type material is referrable to L. angulata as defined here and it seems unnecessary to 
recognize the second species. Picard (19586 : 191) also suggested this synonymy, without 

Laomedea calceolifera (Hincks, 1871) 
(Fig. 18) 

? ] Laomedea exigua Sars, 1857: 50-5 1 . 

Campanularia calceolifera Hincks, 1871 : 78-79, pi. 6; Faure, 1965 : 419^26, figs la, c, 2c, d, 3c, d; 

Miller, 1973 : 377-386, figs 6c, d, 10 a-d. 
Campalaria conferta Hartlaub, 1897 : 495-496, pi. 19, figs 2-12. 
Laomedea angulata: Babic, 1912 : 45 7^60, figs l-5;DaCunha, 1944 : 63, fig. 36. 
Laomedea conferta: Splettstosser, 1924 : 403-420, figs X'-Z 1 , A 2 -T 2 ; Da Cunha, 1944 : 63-64. 
Laomedea calceolifera: Billard, 19316 : 390; Picard, 1955 : 187. 
Obelia calceolifera: Picard, 1955 : 187. 

Laomedea (Campalaria) conferta: Hamond, 1957 : 3 1 5, fig. 23. 
Eulaomedea calceolifera: Rees & Thursfield, 1965 : 102; Millard, 1975 : 223-224, fig. 73g-k. 

TYPE LOCALITY AND MATERIAL. Salcombe, Devon, England, 'on stones &c', offshore (Hincks, 
1871:73-74, 79), several colonies in spirit and one microslide preparation; BMNH 
1899.5.1.155. Some of the spirit material is attached to a worm tube, possibly a sabellariid 
(det. J. D. George). 

TYPE MATERIAL OF OTHER SPECIES. Prof. Dr M. Dzwillo tells me that there is no material 
labelled "Campalaria conferta Hartlaub' in the Zoologisches Institut und Zoologisches 
Museum, University of Hamburg, where Hartlaub worked; and the type material is probably 



therefore lost. The original illustrations were detailed, however, and some appraisal of 
Laomedea conferta is possible. 
I could not locate type material of Laomedea exigua Sars. 

OTHER MATERIAL EXAMINED. All BMNH non-type material is listed. Camara do Lobos, 
Madeira, 146m ('80 fms'), coll. & pres. R. Kirkpatrick, 9 colonies in spirit; 1922.3.4.6. 
'Probably Woods Hole', Massachusetts, U.S.A., ex James Ritchie collection, parts of two 
colonies on microslide; 1964.8.7.83 (mentioned, Rees & Thursfield, 1965 : 102). Cape 
Town, Republic of South Africa, on moored raft, 15 Dec 1949, rf & 9 colonies on two 
microslides; South African Museum coll. SH 423 (Fig. 18). 


Fig. 18 Laomedea calceolifera. (a) part of 9 colony, showing two mature gonothecae and, 
uppermost, an immature one. 'Probably Woods Hole', ex James Ritchie colln (mentioned, Rees 
& Thursfield, 1965 : 102); 1964.8.7.83. (b-c) optical sagittal section and sketch of part of 
lowermost gonotheca in (a), (d) cf gonotheca, Gulf of Pago, NW Yugoslavia; after Babic (1912). 
(e) cf, Cape Town docks, ex South African Museum, microslide preparation SH423. Scale (a-c, 
e, and probably d) 500 /urn. 

DESCRIPTION. Mature colony comprising one to several erect, monosiphonic stems inserted 
at short, irregular intervals on a smooth, tortuous hydrorhiza. Stems probably usually 
unbranched, flexuose; internodes slightly curved to almost straight, long, 3-10 annulations 
basally. Terminal tendrils not recorded. Hydrothecae campanulate, delicate, flared distally, 
length c. \{ x breadth, thin walled, rim even; diaphragm transverse to oblique; pedicels 3-20 
ringed, sometimes with smooth central portion (Fig. 18). Hydranth (BMNH 1964.8.7.83) 


with 15-20 tentacles; hypostome large, probably sub-spherical in life. Gonothecae in axils, 
single or in clusters of up to 3; 9 club-shaped with sub-terminal introverted curving tubular 
aperture on one side, several embryos developing internally; d 1 roughly cylindrical, elongate, 
tapering gradually below, more sharply above, aperture terminal, central, at end of 
introverted tube (as 9 but straight). Young 9 gonotheca truncate (Miller, 1973), lacking 
distinctive aperture until late in development (? young d 1 similar). Reproduction described in 
detail by Miller. No medusa stage. 
Variation. Little information. Male gonothecae vary in width (Fig. 18). 

DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Splettstosser (1924, as C. 
conferta) described the early stages in sporasac development and more recently Miller (1973) 
gave a detailed account of the whole reproductive cycle. Miller interpreted the sporosacs as 
retained medusae ('gonomedusae'). 

REPRODUCTIVE SEASON. Fertile material recorded April-August in NW France (Teissier, 
1965), June off Norfolk (Hamond, 1957), December in the austral seasons of South Africa 
(present material). 

DISTRIBUTION. Widely distributed in warmer parts of the Atlantic Ocean, from South Africa 
(Millard, 1975) north to the English Channel and southern North Sea on the European coast 
and to Maine (Fraser, 1946) on the American side. However, published localities are few. 
For example the only definite British record since Hincks' description of the type material 
is from off Norfolk (Hamond, 1957, as L. conferta). Possibly the several British records of L. 
exigua Sars refer to this species: Swanage, Dorset (Hincks, 1868; possibly BMNH 
1899.5.1.153 is this material); KJlve, N Somerset (Bassindale, 1941); and Great Yarmouth, 
Norfolk (Harmer, in Hamond, 1957). Other European records are as follows: R Ranee and 
Roscoff, NW France (Philbert, 19356; Faure, 1965; Teissier, 1965), Helgoland (Hartlaub, 
1897, as Campalaria conferta and Portugal (Da Cunha, 1944, as L. angulata and L. 
calceoliferd). The species has been recorded from the 'Mediterranean Sea' by several authors 
(Babic, 1912, as L. angulata; Billard, 19316; Philbert, 19356; Picard, 19586) and, more 
precisely, from Castiglione, Algeria (Picard, 1955, as Campanularia conferta). Lastly Linko 
(1911) reported material from Sebastapol, Black Sea. Naumov (1969) stated that the species 
occurred along the 'Atlantic coast of England' and in the North Sea but he confused 
calceolifera with angulata, to which his notes may partly refer (see also Remarks 
concerning 'C. exigua'). 

HABITAT. Extreme lower shore (Faure, 1965) and shallow offshore (Hincks, 1871), 
sometimes deeper. Millard (1975) listed a record of an infertile colony from 70 m off South 
Africa; and one of the specimens listed above was said to come from 146 m ('80 fins') off 

Faure (1965) and Teissier (1965) recorded a usual substrate of rocks and pebbles. 
However, some authors (Hartlaub, 1897; Splettstosser, 1924; Vervoort, \946a; all as 
'conferta') have noted an affinity for Sabellaria tubes (see also notes under Type material) 
while other 'living' substrates recorded include a spider crab (Hyas araneus) carapace and an 
ascidian (Styela coriacea) (both by Hamond, 1957, as Laomedea conferta), and intertidal 
Lithothamnion pools (Teissier, 1965). Morri (1979<a, 6) recorded L. calceolifera in a range of 
salinities down to 27%oin brackish lagoons in Italy. 

REMARKS. For distinguishing features from Laomedea angulata see that species. Infertile 
specimens can be difficult to distinguish from some young, infertile L. angulata and Obelia 
dichotoma colonies. 

Type material of the nominal species Campalaria conferta Hartlaub, 1897, could not be 
located but the original description and detailed illustrations agree with the type material of 
the present species in vegetative features. A difference, however, is that the material 
described by Hartlaub had gonothecae with truncated ends. Miller ( 1 973) has shown that the 
peculiar asymmetric terminal region with its introverted tubular aperture forms late in 9 



gonotheca development, indicating that Hartlaub's account was probably of immature L. 
calceolifera material. Splettstosser (1924) and Hamond (1957) probably also had immature 
material which, following Hartlaub, they assigned to C. conferta. 

The identity of Laomedea exigua Sars, 1857, remains obscure. The species was based on 
Gegenbaur's (1854) material from Sicily briefly described as Campanularia sp. nov. It might 
prove conspecific with calceolifera, which it predates. Van Beneden (1866), Hincks (1868) 
and Leloup (1947, 1952) assigned English Channel material to exigua but it seems likely 
their material was male calceolifera. Naumov (1960, 1969) provided a redescription of L. 
exigua, copying Hincks' figure of a colony and a gonotheca, already copied from Gegenbaur. 
He recorded the nominal species from the Black, Adriatic and Mediterranean Seas and the 
eastern North Atlantic from 'Gibraltar to the North Sea inclusive', but did not cite material. 
Probably his records refer to calceolifera. Bassindale's (1941) record from SW England might 
also have been young L. calceolifera. 

Dimorphic gonothecae were probably first recognized in L. calceolifera by Babic (19 12, as 
L. angulatd), whose illustration of the male gonotheca is redrawn here. 

Stolon growth was studied by Wyttenbach et al. (1973). 

Laomedea flexuosa Alder, 1857 
(Fig. 19) 

Laomedea flexuosa Hincks, in Alder, 18566 : 440 (nom. nud.); Alder, 1857 : 122-123. 
Campanularia flexuosa: Hincks, 1868 : 168-170, pi. 33. 

non Eulaomeda flexuosa: Rees & Thursfield, 1965 : 102-103 (lapsus pro Eulaomedea; redet. Obelia 

Fig. 19 Laomedea flexuosa. (a) New England, USA; 1915.3.6.7. (b) same, rf gonotheca. (c) 9 
gonotheca, ?British Isles: 1912.12.21.290a. (d) part of colony with internodes longer than 
normal, S Greenland; 1 938.3. 1 .297. Scale (a-d) 500 urn. 


NOMENCLATURE. The original publication of the name flexuosa (Hincks, in Alder, 18566) 
lacked description, figures or other definitive indication but a description was published a 
year later (Alder, 1857). The species has been widely recognized and a lengthy synonymy is 

TYPE MATERIAL AND LOCALITY. The type locality has been restricted to the British Isles by 
Cornelius & Garfath (1980). We listed the syntype material, which is preserved in the 
Hancock Museum, Newcastle upon Tyne. 

OTHER MATERIAL EXAMINED. BMNH series, about 50 microslide preparations. 

DESCRIPTION. Colony comprising often branched, erect stems up to c. 30 mm arising at 
irregular intervals from a straight, branching stolon. Stems and branches flexuose; internodes 
characteristically curved, length varied between colonies but constant within, each having 
1-10 annuli basally. Hydrothecae campanulate, robust, sometimes asymmetrically thick- 
ened, length equal to or slightly longer than greatest width; rim even, diaphragm transverse; 
pedicels 3-20 ringed, slightly tapering distally, occasionally with smooth central portion. 
Hydranth with 17-23 tentacles, hypostome spherical. 9 gonotheca subcylindrical, tapering 
basally, truncate distally; sides smooth to slightly sinuous; opening wide, terminal; pedicel 
short, ringed, rf shorter, tapering distally to narrower aperture than in 9. Nematocysts 
described by Ostman (1979). No medusa stage. 

Variation.The following features are variable in the BMNH series: length : breadth ratio and 
radius of curvature of internodes (between, not within, colonies); length of hydrothecal 
pedicel and number of annuli; and gonothecal shape. 

DISPERSIVE STAGE. Planulae, which develop within the gonotheca. Miller (1973) interpreted 
the gonophores as retained medusae ('gonomedusae'). Another description of their develop- 
ment was given by Goette ( 1 907). 

REPRODUCTIVE SEASON. Fertile material recorded April-July, Isle of Man (Bruce, 1948); 
June-July, 1934, Northumbria (H. O. Bull, in Evans, 1978); May-December, NW France 
(Teissier, 1965); February-March, May, September, S Spain (Chas Brinquez & Rodriguez 
Babio, 1977); November, Naples (Lo Bianco, 1909). 

DISTRIBUTION. Recorded widely in the North Atlantic but status in some areas unclear. 
There are records from the Murman and White Sea coasts and Norway (intertidal pools, 
Mathiesen, 1928), the Faeroes and Iceland (Kramp, 1929, 1938), Hardanger Fjord, Norway 
(Brattegard, 1966) and the Shetlands southwards (Hincks, 1868; many other British 
authors). Records from warmer areas are numerous: Santander, N Spain (Rioja y Martin, 
1906), S Spain (Chas Brinquez & Rodriguez Babio, 1977), Portugal (Da Cunha, 1950), 
Mauritania (Billard, 1906), Morocco (Patriti, 1970), Ghana (Buchanan, 1957); but not 
southern Africa (Millard, 1975). In the Mediterranean Sea the species has been recorded 
from Naples (Lo Bianco, 1 909; Riedl, 1 959) and 'E Adriatic' (Pieper, 1 884). 

The northern and southern limits on the North American coastline are also unclear. 
Although Kramp (1938) reported the species from W Greenland, Calder (1970) did not 
record it from Canadian waters and Eraser's ( 1 944) most northerly record was from the Gulf 
of St Lawrence. The species has been reported along most of the United States coast, but the 
status in the Caribbean is just a single record which was doubted by both Fraser ( 1 944) and 

HABITAT. Recorded on a wide variety of inert and living substrates (Hincks, 1868; Vervoort, 
1946a; Marine Biological Association, 1957; Teissier, 1975; Chas Brinquez & Rodriguez 
Babio, 1977), probably lacking a strong substrate preference although often found on fucoid 
algae. Occurs intertidally, and probably not often deeper than the 37 m recorded by Miller 
( 1 96 1 ) and the 40 m from which a BMNH specimen was collected (reg. no. 1971.5.11 .24, W 
Scotland). However, a deep record from '12 positions 20^48 miles (32-77 km) SW of 
Eddystone', SW England, 73-92 m (Marine Biological Association, 1957), suggests the 
species is not unusual at such depths; and Crawshay (1912) reported the species from 


80- 100m at several stations in the western English Channel. The four microslide 
preparations listed under this species by Rees & Thursfield (1965), from depths down to 
160 m, are ofObelia dichotoma material. 

REMARKS. Laomedea flexuosa is type species of the unacceptable genus Eulaomedea Broch, 
1910, and of the genus Laomedea (discussion of both on pp. 97-98). 

Alder (1856/7) introduced the name flexuosa in passing when describing Laomedea 
neglecta in the phrase 'L. flexuosa Hincks, MS'; but the introduction was invalid as it lacked 
an acceptable indication. It is clear that in or shortly before 1856 Alder and Thomas Hincks 
concurred thai flexuosa was a valid species, and were the earliest to recognize it. When first 
mentioning the name Alder (18566) cited as indication 'the young of Johnston's small 
variety of L. gelatinosa'; but Johnston (1847 : 472) said that he had not seen material. He 
stated simply: 'variety a with even margins and simple stem (not seen by me)'. Johnston 
apparently referred to hints by earlier authors that there was a species awaiting 
recognition. But Johnston did not see flexuosa material himself, nor did he cite this earlier 
literature. Hence Alder's (18566) indication of Johnston's brief account does not satisfy the 
requirements of nomenclature, and the name flexuosa must date from the later paper (Alder, 

The earlier mentions of the species which Johnston (1847) apparently had in mind are 
not easily traced. He may have thought that Couch (1844 : 39-40) had seen some flexuosa 
material when, in describing gelatinosa, he stated that species often to be just one inch 
(2 5 '4 mm) high compared with the greater height usual in gelatinosa s. str. It can be noted 
that Couch might have included neglecta also within gelatinosa, since he recorded that 
'gelatinosa' sometimes grew under stones. Following Couch's early death it was left to Alder 
and Hincks to define correctly the three species involved. Thus, Alder (18566 : 440) stated 
that 'if observed [by earlier workers possibly Couch] neglecta has been passed over as ... L. 

L. flexuosa has been used by several experimental biologists. Stolon growth has been much 
studied and was mentioned above (p. 42). Phenotypic response to simple environmental 
changes was described by Crowell (1957; 1961; see p. 42). Embryonic rupture of the 
hydrothecal bud was studied by Knight (1965, 1970, 1971). Stebbing (1976, 1979) investi- 
gated the influence of inorganic toxins on growth, noting that sub-inhibitory doses actually 
enhanced growth rates. Factors such as straightness of stolon, growth rate and frequency of 
gonophore production were all affected by water quality. In a later paper (Stebbing, 1981) he 
studied several factors affecting growth. 

The material assigned to this species by Rees & Thursfield (1965) I refer to Obelia 

Laomedea neglecta Alder, 18566 
(Fig. 20) 

Laomedea neglecta Alder, 18566 : 440, pi. 16, figs 1-2; Vervoort, \946a : 308-310, fig. 316 (syn. L. 

brochi Splettstosser; L. lauta Hummelinck); (non DaCunha, 1950 : 142-143, fig. 9). 
Campanularia fragilis Hincks, 1863 : 46-47; Hincks, 1868 : 175-176, pi. 32, fig. 3 (syn. C. elongata 

Van Beneden). 

Laomedea decipiens Wright, 1863 : 49, pi. 5, fig. 9. 

Campanularia elongata Van Beneden, 1866 : 164-165, fig. 6 (p. 150); Leloup, 1947 : 24-25. 
Campanularia neglecta: Hincks, 1868: 171-172, pi. 30, fig. 2; Hincks, 1872 : 390-391, pi. 20, fig. 4. 
Campanularia decipiens: Hincks, 1868: 173-174. 
Laomedea brochi Splettstosser, 1924 : 376-403, figs A-Z, A -W.' 
Laomedea ?lauta Hummelinck, 1930 : 35-37, fig. 5. 
Laomedea (Paralaomedea) neglecta: Hummelinck, 1936 : 57 (syn. ?L. brochi Splettstosser; L. lauta 


TYPE MATERIAL AND LOCALITY. Single infertile colony preserved as two microslide 
preparations, Hancock Museum, Newcastle upon Tyne, Northumberland; designated 


neotype by Cornelius & Garfath (1980). Although the neotype was once part of Alder's 
collections and was probably identified by him it almost certainly came from an Essex 
locality, whereas most of Alder's collections came from NE England. The type locality 
remains 'Cullercoats and Tynemouth, Northumberland, on undersides of stones' (designated 
by Nutting, 1915; repeated by Cornelius & Garfath). 

TYPE MATERIAL OF OTHER SPECIES EXAMINED. Laomedeo. lauta Hummelinck, 1930, holotype; 
Rijksmuseum van Natuurlijke Historic, Leiden, reg. no. 259. Hummelinck's illustration of 
this material is very accurate. 

OTHER MATERIAL EXAMINED (L. neglecta is only poorly represented in the BMNH collec- 
tion). Forth Cressa, St Mary's, Isles of Scilly, on underside of block of granite, c. LWM, 17 
Apr 1903, fertile colonies, 'ova in an external capsule' (ms note by E. T. Browne, Zoological 
Notebook 10 : 15, preserved in BMNH), spirit + 2 microslide preparations, coll. E. T. 
Browne; 1948.9.8.1 19 (Fig. 20). Mewstone Echinoderm Ground, near Plymouth, Devon, c. 
50 m, October 1899, infertile colonies, coll. Marine Biological Association of the U.K., det. 
E. T. Browne (ms note in Zoological Notebook 13 : 149), spirit material (1948.10.1.126)+ 1 
microslide preparation (1959.9.17.19). Gaso Ranna, Gullmarsfjord, Sweden, 20-30 m, 27 
Aug 1962, infertile colonies, spirit material + 1 microslide preparation, coll. W. J. Rees; 
1962.11.8.9. Zooligisch Museum, Amsterdam: Brehorn, Zuider Zee, Netherlands, 14 Jun 
1927, infertile and <5 fragments in spirit (Zuider Zee Onderzoek sta. 1; det. & mentioned 
Hummelinck, 1936 : 57, fig. 9 a-g) (Fig. 20). Dollard, NE Netherlands, fertile colony in 
spirit, coll. A. P. C. de Vos, 9 Aug 1954. Kornwenderland, NE Zuider Zee, 4-6 m, coll. 
Zoological Station, den Helder, 29 Sep 1938, several infertile fragments in spirit, det. W. 
Vervoort. Rijksmuseum van Natuurlijke Historie, Leiden: Aberystwyth Bay, Wales, 1 7 Jun 
1939, intertidal, several infertile fragments, coll. P. W. Hummelinck; RMNH reg. no. 968. 
Strand Renesse, Schouwen, Netherlands, 20 Dec 1941, infertile fragments on Tubularia 
?indivisa stems, coll. J. Viergever; RMNH reg. no. 702 (some of these specimens are 
polysiphonic, having two hydrocauli fused basally). Kornwenderland, 6m, 29 Sep 1938, 
four small colonies on one microslide, coll. Zoological Station, den Helder, det. W. 
Vervoort; RMNH reg. no. 1012. 

DESCRIPTION. A small species. Hydrorhiza smooth, tortuous, sometimes branched; 
short, delicate monosiphonic to bisiphonic stems arising at irregular intervals. Internodes 
long, narrow, often wider in middle than at ends, 3-10 rings basally, curved (recalling L. 
flexuosa) to almost straight (e.g. Hummelinck, 1936 : fig. 9a); each sharply inturned basally; 
hydrothecal pedicel on short distal process in line with axis of internode. Hydrotheca long, 
delicate, unthickened, cylindrical; length = 3 x width; diaphragm oblique to transverse; rim 
usually bimucronate but often abrades smooth; pedicel long, tapering distally, up to c. 20 
annuli, sometimes with 1-3 smooth central portions. Hydranth with c. 20 tentacles (Van 
Beneden, 1866, as C. elongata; present material, 1971.5.11.11). Gonotheca (after 
Splettstosser, 1924, as L. brochi) rf = 9, cylindrical to inverted-conical, truncated above. 
Acrocyst in 9, eggs possibly extruded singly (Hincks, 1868; Splettstosser, 1924). 
Variation. The delicate hydrothecal rim easily abrades smooth. Some published descriptions 
suggest that there are minute spines associated with the cusps on the hydrothecal rim but 
these seem simply to be folds. Splettstosser (1924 : fig. B, as L. brochi) illustrated a 
hydrotheca in which the bimucronate condition was obscure, and quite apart from damage 
by abrasion it seems that not all specimens are perfectly bimucronate. Other bimucronate 
species vary in the same way (Obelia bidentata, Clytia paulensis; see also p. 40). 

DISPERSIVE STAGE. Planulae, brooded in 9 acrocyst derived from vestigial retained medusa 
(Splettstosser, 1924). Developing eggs have been reported in the endoderm tissues of the stem 

Hincks (1872) described what was apparently a vegetative spore developing in place of a 
hydranth, on a specimen collected from British waters in June. 



REPRODUCTIVE SEASON. Probably April-October, but infertile colonies not unusual during 
this period. All available information is given: fertile material recorded Netherlands, 14 
June, 1927 (Hummelinck, 1936; also present material) and 9 August, 1954 (present 
material); Isle of Man, August (Bruce et al., 1963); Norfolk, England, 18 September, 1954 
(Hamond, 1957); Northumbria, June (J. H. Robson, in Evans, 1978); SE Scotland, October 
(Laverack & Blackler, 1974); Scilly Isles, 27 April, 1903 (present material). 

Infertile material recorded Netherlands, 27 August, 1929 (Hummelinck, 1936), 29 
September, 1938 & 20 December, 1941 (present material); Aberystwyth, Wales, 17 June, 
1939 (present material); Norfolk, 2 July, 1952 (Hamond, 1957); W Sweden, 27 August, 
1962; SW England, October, 1899 (present material). 

Fig. 20 Laomedea neglecta. (a) part of colony, Zuiderzee, Cl = 14'8%o; Amsterdam Zoological 
Museum collection, (b) same, one hydrotheca. (c) same, part of hydrothecal rim. (d) gonotheca 
with acrocyst, Is of Scilly, SW England; 1948.9.8. 1 19. Scales: (a, c-d) 500 ^m; (b) 100 //m. 

DISTRIBUTION. Although infrequently reported L. neglecta seems widely distributed in the 
North Atlantic. The most northerly and southerly records are from Iceland (Kramp, 1938) 
and the Adriatic Sea (Vatova, 1928). All other published records are listed (see also Material 
list): England (Alder, 1 856/7; Hincks, 1868, 1872; Hamond, 1957); Isle of Man (Herdman, in 
Wood, 1901 : 20; Bruce et a/., 1963, as Campanularia fragilis); Scilly Isles (Vallentin, in 
Browne & Vallentin, 1904; repeated in Robins, 1969); SE Scotland (Wright, 1859, as 
Laomedea decipiens, see Remarks; Laverack & Blackler, 1974); Aberystwyth, Wales 
(present material); Netherlands (Vervoort, 1946a); Belguim (as C. fragilis, Van Beneden, 
1866, repeated in Leloup, 1947, 1952); Kattegat (Jagerskiold, 1971; Rasmussen, 1973); W 
France (Billard, 1927). 

Picard (19586) did not include the species in the mediterranean faunal list but Vatova's 
(1928) adriatic record seems soundly based. The records by Broch (1933), from the Adriatic, 


and Da Cunha (1950), from Portugal, were probably based on Obelia bidentata material. 
L. neglecta has also been recorded from the eastern coast of North America (Fraser, 1944). 

HABITAT. Intertidal to at least 50 m. All available information is given: intertidal, under- 
neath stones and on other hydroids (in Britain, Alder, 18566; Hincks, 1868); under stones, 
on Tubularia stems and at 50 m depth (present material); 1 5-50 m on sea-beds of mud, clay 
and shell gravel (Kattegat, Jagerskiold, 1971); 'on stone embedded in meshes of whelk pot', 
offshore, and under rocks in tidal creek (SE England, Hamond, 1957); on pebbles & oysters 
(W France, Billard, 1927); LWM, underside of rock (Scilly Isles, present material, Browne & 
Vallentin, 1904; repeated in Robins, 1969). 

Collected in salinity of 26'7%ofrom Zuider Zee (Hummelinck, 1936 : 57; data with speci- 
men in Zoologisch Museum, Amsterdam). 

REMARKS (see also Remarks under Laomedea flexuosa). Laomedea neglecta is a small 
species and although widespread has been little recorded. It was first described from English 
waters but there are still only a few records from Britain and Europe. 

Infertile colonies resemble Obelia bidentata, and there is some overlap in hydrothecal 
length. Although the stems of L. neglecta are probably always narrower than those of O. 
bidentata, identification of small specimens can still be difficult. 

The spines associated with the hydrothecal rim by several authors are merely folds in the 
hydrothecal wall appearing as artefacts during preservation. 

Campanularia fragilis Hincks, 1863, was probably founded on a colony of the present 
species. The original illustration was actually published, without a binominal, a year earlier 
than the species name and description, in vol. 10 of the same journal, forming plate 9, figure 
3. The illustration seems to represent L. neglecta. Hincks distinguished the two species on 
the 'markedly flexuose character of the stem', the long hydrotheca with plain rim and the 
small size of the colony. Probably the hydrothecal rims had abraded smooth. The holotype 
(not found) was collected from under stones in a rock pool, a likely habitat for L. neglecta. 
Hincks later (1868) referred to the species as C. ?fragilis, and significantly regarded C. 
elongata Van Beneden, 1866, type locality Ostend, Belgium, as conspecific. Vervoort 
(1946a :310) quoted Maitland's (1897) opinion that elongata and fragilis were conspecific; 
an opinion Vervoort considered to have been a pers. comm. from Van Beneden to Maitland. 
Leloup (1947) concurred with this synonymy. There seems to have been no material 
recorded as elongata since the original description. Van Beneden illustrated a long 
hydrotheca with even rim and described the (infertile) colony as small. The hydranth had c. 
20 tentacles. His description, like that of Hincks, probably represents L. neglecta with 
abraded hydrothecal rims. 

Laomedea decipiens Wright, 1863, although described with some precision, was 
illustrated only by a small woodcut of a hydrotheca. Wright likened the species to L. neglecta 
except that the hydrothecal rim was 'even, and had the appearance of being double for about 
half its length from the rim', arguably a misinterpretation; and that 'the reproduction' of L. 
decipiens resembled 'exactly' the process he had described (Wright, 1859) in Opercularella 
lacerata (Johnston, 1847). That is, there was an external acrocyst. But Wright said his 
'decipiens' material differed in that each acrocyst contained only three ova, while those of 0. 
lacerata had seven or eight. L. neglecta is now well known to have an acrocyst containing 
rather few ova. Further, its hydrothecal rims often wear smooth. Hence it is plausible that 
decipiens was based on fertile material of neglecta in when the hydrothecal rims had worn. 
Hincks (1868) repeated Wright's description and indicated that Wright had obtained his 
material from the Firth of Forth Wright had given no locality. Wright and Hincks were in 
close contact (Hincks, 1868 : preface) and Wright probably told Hincks the locality. Hincks 
did not know the gonosome of neglecta and was rightly cautious when treating decipiens. 
The record from the Firth of Forth was repeated uncritically by Leslie & Herdman (1881) 
and Pennington ( 1 885), but the species seems not to have been recognized since. 

Laomedea brochi Splettstosser, 1924, and Laomedea lauta Hummelinck, 1930, were 



referred to the present species by Hummelinck (1936) and Vervoort (1946a) with good 
reason. See also the notes under Variation. 

Da Cunha (1950) recorded L. neglecta from Portugal growing on a species of Sertularella. 
However, his illustration shows a medusa developing within a gonotheca and his material 
was probably Obelia bidentata. 

The affinities of L. neglecta are discussed above (p. 47-49). 

Laomedea pseudodichotoma Vervoort, 1959 
(Fig. 21) 

Laomedea (Eulaomedea) pseudodichotoma Vervoort, 1959:316-318, figs 56-57; Vervoort, 
1966: 104. 

TYPE MATERIAL AND LOCALITY. Lectotype designated from original syntype series by 
Vervoort (1966), 50mm d 1 colony with developing and mature gonothecae, 1343'N, 
17 23' W (off Senegal); mostly preserved in University Zoological Museum, Copenhagen, 
with part on microslide in Rijksmuseum van Natuurlijke Historic, Leiden. 'Paralectotypes', 
9 colonies, from 5 37' N, 38' E (off Ghana); also preserved in Copenhagen. 

OTHER MATERIAL EXAMINED. Off Abidjan, Ivory Coast, 35m, 2 Mar 1966; Rijksmuseum 
van Natuurlijke Historic, Leiden, reg. no. 10410 (Fig. 21). 

DESCRIPTION (partly after Vervoort, 1959, 1966). Colonies so far recorded up to 50mm, 
polysiphonic basally; stem almost straight; roughly pinnate; some secondary branching; 
branches approximately alternate, in one plane, in the only available colony. Internodes 
slender, 2-3 ringed basally; some tanning. Hydrothecal pedicels short, 5-10 ringed; some 
axillary. Hydrothecae delicate, conical, some slightly swollen below; diaphragm oblique in 
side view, basal chamber large; rim even, circular. Gonothecae dimorphic, d 1 long, 
cylindrical to slightly tapering basally, widest about from aperture; truncate, aperture 
simple, not raised, as wide as gonotheca. Immature cf gonotheca much shorter. 9 gonotheca 
long, widest in terminal |, tapering gradually below and abruptly above; truncate; aperture 
raised, half width of gonotheca; gonophores thought to be heteromedusoid, sessile. 

Fig. 21 Laomedea pseudodichotoma. (a-c) part of colony, single hydrotheca and 9 gonotheca, 
Abidjan, Ivory Coast, 35 m. (d) rf gonotheca, off Senegal; syntype series. Scales: (a) 500 //m; 


DISPERSIVE STAGE. Almost certainly no medusa (Vervoort, 1959). Presumably planulae are 
released in the normal way. 

REPRODUCTIVE SEASON. Fertile material recorded 30 January, 1946, 50 m, off Ghana, 9; 2 
March, 1966, 35 m, off Ivory Coast, 9; 25 April, 1946, 65-89 m, off Senegal, rf; (Vervoort, 
1959; present material). 

DISTRIBUTION. Recorded only from the coastal waters of Ghana, Ivory Coast and Senegal. 

HABITAT. So far found only on Sertularella cylindritheca (Allman, 1888), at depths from 
35 m to 89 m (Vervoort, 1959; present material). 

REMARKS. This species is known only from tropical west Africa. 

Genus OBELI A Peron & Lesueur, 

Obelia Peron & Lesueur, 18100 : 355; Peron & Lesueur, 718106 : 43. 

Monopyxis Ehrenberg, 18340 : 297; Gray, 1848 : 84. 

Obelomma Haeckel, 1879 : 176. 

Obeliopsis Le Danois, 1913:110. 

For other synonymy see Remarks, Cornelius (\915a) and page 114. See also note 3 (p. 124) 

concerning the date of introduction of the genus name Obelia. 

TYPE SPECIES. Obelia sphaerulina Peron & Lesueur, 1810# (nom. nov. pro Medusa marina 
Slabber, 1769); by monotypy. For taxonomic purposes the type species was taken to be 
conspecific with hydroid O. dichotoma (Linnaeus, 1 758) by van der Hoeven ( 1 862 : 280) and 
Russell (1953 : 297), but this link is subjective. For the time being at least there is great 
difficulty in relating medusae collected from the plankton to their hydroids (e.g. Cornelius, 
1975#). Peron & Lesueur cited as indication a specimen taken in Dutch waters, as illustrated 
in the German edition of Slabber's (1775-1781 : pi. 9, figs 5-8) work. But Goy (1980 : 72) 
links also to the published description an unpublished illustration by Lesueur. The specimen 
illustrated therein was taken near Le Havre, in the Museum of which town the illustration is 
preserved (see also note 3, p. 124). 

Naumov (1960, 1969) and Stepanyants (1979) designated Sertularia geniculata Linnaeus, 
1758, type species of Obelia; but geniculata was not an originally included species. For 
discussion see Cornelius ( 1 91 5a, 1981). 

DIAGNOSIS. Colonial Campanulariidae with: polyp generation forming upright colonies, 
branched or unbranched, variably flexuose; stolon not anastomosing; internodes annulated 
proximally, supporting hydrotheca on distal lateral process; hydrotheca bell shaped, 
hydranth with sub-spherical hypostome; gonotheca inverted cone-shaped, usually with 
raised terminal aperture but sometimes simply truncate; medusa umbrella flat, eversible, 
mesoglea thin; velum reduced to absent; manubrium long; about 16 marginal tentacles on 
release, numerous in adult, short. 

REMARKS. An exhaustive synonymy and restriction of this genus have already been 
published (Cornelius, 1975a) but the most recent redefinitions of Obelia are by Stepanyants 
(1979) and Arai & Brinkmann-Voss (1980). The genera referred to Obelia by Cornelius 
included: Slabberia Oken, 1815 (a 'rejected work' for nomenclatural purposes); 
Campanularia Lamarck, 1816 (part; but see p. 51); Thaumantias Forbes, 1848 (part; see 
also p. 71); Eucope Gegenbaur, 1856 (part; here referred to Clytia, see p. 71); 
Schizocladium Allman, 1871; Obelaria Haeckel, 1879 (but see p. 94); Obeletta Haeckel, 
1879; Obelissa Haeckel, 1879; and Monosklera von Lendenfeld, 1885. The nomenclatural 
problems involving Obelia with Laomedea are discussed above (p. 97). 

Three species of Obelia are recognized from the hydroid stage in the eastern North 
Atlantic but their medusae cannot be told apart (Russell, 1953; Cornelius, 1975a; Arai & 
Brinckmann-Voss, 1980). Two of the species were described from the hydroid stage before 


any of the medusae, and their 'hydroid names' can be regarded as having safe seniority. The 
description of the third valid hydroid stage, O. bidentata Clarke, 1875, post-dates several 
binominals applied to the medusa stage and this name cannot yet be regarded safe. But there 
are difficulties in identifying to species level the early descriptions of the medusa and linking 
them with the correct hydroid stages. Thus it seems probable that the name O. bidentata 
will be unchallenged for some time, and that a working stability has been reached. 

The subgenus Monopyxis was introduced in the combination Sertularia (Monopyxis) 
geniculata Linnaeus, 1758, by Ehrenberg (1834a : 297). S. geniculata is type species of the 
subgenus, by monotypy. Since there are only 3-5 species of Obelia recognized or 
provisionally accepted from the hydroid stage (e.g. Cornelius, \915a) it seems superfluous to 
subdivide the genus, and Monopyxis can be referred to Obelia. Ehrenberg's account was 
based on material from Norway, Denmark and Germany. Gray (1848 : 84) used Monopyxis 
as a supergenus, including in it the species Obelia geniculata, Hartlaubella gelatinosa, 
Campanularia flemingii (now referred to H. gelatinosa, p. 96) and Monotheca obliqua (e.g. 
Hincks, 1868, as Plumularia); but this supergenus name was not employed again. Finally 
Hincks (1868) included Monopyxis in his synonymy of Obelia. 

The genus Obelomma Haeckel, 1879, was established to include Obelia medusae with 48 
tentacles on release. It included three nominal species, each incorporating several others 
within the synonymies Haeckel presented. Most of those incorporated are now referred to 
Obelia, of which Obelomma should be regarded a synonym. 

The generic names Obelaria Haeckel and Obelaria Hartlaub are considered under 
Hartlaubella (p. 94); and Eucope Gegenbaur is treated under Clytia (p. 7 1 ). 

The genus Obeliopsis Le Danois, 1913, was erected to include material referred to a single 
species (see p. 120) and can be confidently referred to Obelia. 

General notes on the medusa stage and indications to other literature are given under O. 
dichotoma (p. 118) and in Cornelius (1975<a). Applications of the genus name Obelia to 
bryozoan species were also listed in that paper. 

The three species of Obelia (hydroid) occurring in the NE Atlantic can usually be 
identified from the characters given by Cornelius (1975a : table 1); but occasional specimens 
of O. dichotoma approach O. geniculata in having a slight thickening of the internodal 
perisarc. Some of these specimens cannot be confidently identified to species. 

Obelia bidentata Clarke, 1875 
(For illustrations see Cornelius, \915a) 

Obelia bicuspidata Clarke, 1875 : 58, pi. 9, fig. 1; Stepanyants, 1979 : 37-38, pi. 7, fig. 1 (syn. O. 

austrogeorgiae Jaderholm, 1904a; Clvtiapaulensis Vanhoffen, 1910). 
Obelia bidentata Clarke, 1875 : 58-59^ pi. 9, fig. 2; Jaderholm, 19046 : 270-271 (syn. O. bicuspidata 

Clarke; first reviser); Cornelius, 1975a : 260-265, fig. 2 (syn. O. bicuspidata Clarke). 
Clytia arborescens: Billard, 1 907 : 1 67 (non Pictet, 1 893). 
For further synonymy see Cornelius ( 1 975a) and Table 4. 

NOMENCLATURE. Jaderholm (19046) acted as first reviser when using the specific name 
bidentata for this species, and several contemporary authors followed him (references in 
Cornelius, \975a). Although the superseded name bicuspidata has been widely used, O. 
bidentata is the available name under the current conventions of zoological nomenclature. 

DISPERSIVE STAGE. The medusa of this species has probably still to be reared to maturity but 
when young resembles those of the other two Obelia species (Cornelius, 1975<2, \911a). I did 
not mention in the 1975# paper that Professor K. Ramunni 'reared the medusa of this 
species' from Bengal material (mentioned in Annandale, 1915, as O. spinulosd). The age to 
which he reared it was not recorded. Ramunni was by far the earliest to realize that the 
medusa resembled that of the other two Obelia species, and to see it released. 

DISTRIBUTION. Recorded from the English Channel and southern North Sea southwards 
through most European and African coastal waters but not from the Black Sea or the Baltic. 



Table 4 Synonymies among the nominal species of Obelia described from the hydroid stage (after 
Cornelius 1975a, with additions). Although the lists are World-wide most of the nominal species have 
been recorded from the eastern North Atlantic. Many were first described in genera other than Obelia. 
Discussion of species treated in the 1975a paper is not duplicated here. The references cited in the 
table are mainly as in that paper, only the few additional ones being included in the present reference 

(a) Synonyms of Obelia bidentata Clarke, 1875 
O. bicuspidata Clarke, 1 875 
Campanularia spinulosa Bale, 1888 

O. andersoni Hincks, 1889 

O. bifurca Hincks, 1889 

Gonothyraea longicyatha Thornely, 1899 (non 

O. longicyatha Allman, 1877) 
O. corona Torrey, 1 904 
Obelia sp. Clarke, 1907 
O. bifurcata Thornely, 1908 
O. multidentata Fraser, 1914 
O. oxydentata Stechow, 1914 
O. longa Stechow, 1921 
Clytia longitheca Hargitt, 1924 
O. longitheca Hargitt, 1 924 (sic) 
O. attenuata Hargitt, 1924 
Laomedea bicuspidata var. picteti Leloup, 1932 
L. spinulosa var. minor Leloup, 1932 
L. bicuspidata var. tenuis Vervoort, 1946 
IClytia arborescens: Billard, 1893 (see present 

paper, p. 1 17) 

(b) Synonyms of Obelia dichotoma (Linnaeus, 

Sertularia longissima Pallas, 1 766 

Sertolare genicolata Cavolini, 1785 (lapsus 

pro Sertularia geniculata) 
ICymodocea simplex Lamouroux, 1816 
ITubularia clytioides Lamouroux, in Freycinet, 


Campanularia maior Meyen, 1 834 
C. brasiliensis Meyen, 1834 
C. cavolinii Deshayes & Edwards, 1 836 
C. caw//m Chiaje, 1841 
Laomedea gracilis Dana, 1 846 
O. commissuralis McCrady, 1857 
L. divaricata McCrady, 1857 
Eucope parasitica Agassiz, 1865 
E. pyriformis Agassiz, 1 865 
E. articulata Agassiz, 1865 
Campanularia flabellata Hincks, 1 866 
O. plicata Hincks, 1 868 
Schizocladium ramosum Allman, 187 1 
O. pygmaea Coughtrey, 1 876 
O. hyalina Clarke, 1879 
O. adelungi Hartlaub, 1884 
O. helgolandica Hartlaub, 1884 
O. australis von Lendenfeld, 1885 
C.cheloniae Allman, 1888 
O. angulosa Bale, 1888 
O. chinensis Marktanner-Turneretscher, 1890 
O. arruensis Marktanner-Turneretscher, 1 890 
O. nigrocaulus Hilgendorf, 1898 
O. gracilis Calkins, 1899 

(b) Synonyms of Obelia dichotoma (Linnaeus, 
1758) (continued) 

O. surcularis Calkins, 1899 

O. fragilis Calkins, 1 899 

O.griffini Calkins, 1899 

O. rhunicola Billard, 1901 

O. borealis Nutting, 1901 

O. dubia Nutting, 1901 

O. solowetzkiana Schydlowsky, 1902 

C. obtusidens Jaderholm, 1905a 

L. congdoni Hargitt, 1909 

O. piriformis Bedot, 1910 

O. pyriformis: Mayer, 1910 

L. sargassi Broch, 1913 

O. undotheca Stechow, 1923 

O. nodosa Bale, 1924 

O. coughtreyi Bale, 1 924 

O. obtusidentata Bedot, 1925 

O. dischotoma Billard, 1927 (lapsus pro 


O. everta Hargitt, 1927 
1O. commensuralis Gudger, 1937 (?lapsus pro 

O. alternata Fraser, 1938 
O. equilateralis Fraser, 1938 
O. microtheca Fraser, 1938 
O. tenuis Fraser, 1938 
O. racemosa Fraser, 1 94 1 
O. irregularis Fraser, 1943 
O. braziliensisVannucci Mendes, 1946 
O. biserialis Fraser, 1 948 
O. hyaliana Vannucci, 1955 (see present paper, 

p. 119) 

(c) Synonyms of Obelia geniculata (Linnaeus, 

Laomedea lairii Lamouroux, 1 82 1 
Campanularia prolifera Meyen, 1834- 
Eucope diaphana L. Agassiz, 1 862 
E. alternata A. Agassiz, 1865 (nom. nov. pro E. 
diaphana L. Agassiz; non Thaumantias 
diaphana A. Agassiz) 
E. polygena A. Agassiz, 1 865 
E. fusiformis A. Agassiz, 1865 
O. gymnopthalma Spagnolini, 1 87 1 
Monosklerapusilla\on Lendenfeld, 1885 
O. geniculata vars /, // & /// Marktanner- 
Turneretscher, 1890 

Campanularia coruscans Schneider, 1 897 
O. geniculata f. subsessilis Jaderholm, 19506 
O. geniculata f. gaussi Vanhoffen, 1910 
O. geniculata ff. intermedia, subantarctica & 
subtropica Ralph, 1956 


There are few English records but from the north coast of France southwards there are many: 
NW France, Belgium, Netherlands (earliest records in Table 5; also Vervoort, \946a; 
Leloup, 1947; Teissier, 1965); Cadiz, Gibraltar, Morocco, Mauritania, S of Madeira 
(Stechow, 1925); Algeria, Senegal (Picard, 19516, 1955); Ghana (Buchanan, 1957); Gambia 
(Vervoort, 1959); Morocco (Patriti, 1970); Azores, Nigeria, Sierra Leone (Cornelius, \915a); 
South Africa (Millard, 1975). The recorded northern limits are north of the Wash (Table 5) 
and Schiermonnikoog in the Frisian Islands (53 30' N, 615'E; RMNH Leiden, W. 
Vervoort, pers. comm.). By coincidence the most southerly records, from South Georgia and 
Tierra del Fuego (Cornelius, \915a), are from a similar latitude, 54 S. A record from Adelie 
Land, Antarctica, may be based on Clytia paulensis material (see Remarks). 

My statement (Cornelius, 1975#) that Kramp (1961) recorded the species from Helgoland 
was incorrect and there are no records from that island (W. Werner, pers. comm.) or from 
any other German waters. The Kent material which I listed was wrongly identified. 

The Dutch record of Leloup (1933 : 1 1) listed in Table 5 was referred by Hummelinck 
(1936 : 56) and Vervoort (19460 : 300) to Hartlaubella gelatinosa; but Leloup also recorded 
gelatinosa in his paper. I have not located Leloup's material but if O. bidentata it would be 
among the earliest Dutch records. 

Some aspects of the European distribution deserve comment. Although the species is 
distinctive and occurs widely in warmer parts of the Atlantic and Indo-Pacific oceans, it was 
not recorded from the eastern side of the Atlantic until the 1900s. It might previously have 
been confused with H. gelatinosa, as suggested by both Billard and Leloup in various 
publications. Confusion certainly occurred in some of the published records of the species 
from Belgium and England (possibly Leloup, 1933; Robins, 1969; both as H. gelatinosa; see 
previous paragraph and Table 5). Further, the species was poorly documented before 1900 
and the descriptions available then were in obscure literature. It is understandable that when 
Billard (1902) recorded the first French material he referred it to Obelia bifurca Hincks, 
1889, described from the Indian Ocean. (O. bifurca is now regarded conspecific.) Much later 
Stechow (1925) too referred material to O. bifurca, 50 years after O. bidentata had first been 
described; and it is evident that news of bidentata had spread slowly. 

Although the European records (Table 5) appear to show the arrival of the species in 
Cherbourg and Ostend in 1902, the Netherlands in 1926, Norfolk in 1951, the Scilly Isles in 
1966 and Hampshire in 1975 (an unusually hot summer), there is no satisfactory proof of a 
gradual extension of range. Indeed, the apparent spread of the species along the coasts of 
France, Belgium and the Netherlands follows quite closely the coming of hydroid experts in 
those countries, by coincidence from west to east (Billard, Leloup, Hummelinck, Vervoort). 
South-east England has seldom been given close attention by hydroid workers. Hamond 
(1957) recorded O. bidentata off Norfolk in 1951 as 'rather common' over a wide area, 
suggesting that it was already long established but for how long cannot be decided. Parallel 
evidence comes from another overlooked species, similarly told from its near relatives by a 
bimucronate hydrothecal rim: Clytia paulensis. Although some BMNH material of 
paulensis was collected from near Plymouth in 1899 (p. 90) it was not identified for 80 
years; and the species was first recognized from British waters independently in 1973. It 
seems that for a long time it too was overlooked and thought to occur no further north than 
NW France; but now it has been found again in southern England and, like O. bidentata, in 
East Anglia. 

It remains puzzling that the nineteenth century English and Belgian experts active around 
the English Channel (Couch, Van Beneden, Gosse, Hincks) did not find O. bidentata. Hincks 
(1889) in fact described the species from Indian Ocean material, as O. bifurca, and would 
surely have recognized British examples. So it might really have been absent at that time; and 
it may be relevant that in 1937-1938 Vervoort (19466) did not record the species from the 
Waddenzee, N Netherlands, where it now occurs. Finally, although O. bidentata has been 
recorded from the well worked offshore region around Roscoff (Billard, 1912; Teissier, 1965) 
it is scarce and has been found only in deep water (L. Cabioch, pers. comm.). See also page 




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HABITAT. Substrates not recorded in my \915a paper include an isopod, Anilocra physodes 
(Linnaeus), from Naples (Stechow, 19216, 1925, as Obelia bifurca) and a commercial 
shrimp, Pennaeus aztecus Ives, from Florida (Kruse, 1959; via A. A. Fincham, pers. 
comm.); ships hulls, hermit crabs gastropod shells and algae (Millard, 1975). O. bidentata is 
highly tolerant of brackish water. It has been recorded at salinities of 18'6%o (Netherlands; 
Vervoort, 19460), 10-20%o (Italy; Morri, 19796) and even so low as l%o (S Carolina, U.S.A.; 
Calder, 1976). The species has often been recorded in shallow water and Millard (1978) 
listed it as intertidal. 

REMARKS. A detailed, world-wide treatment has appeared recently (Cornelius, \915a) and 
the notes given here are mainly intended to supplement that account. O. bidentata is a nearly 
cosmopolitan species, and a possible explanation is given above (pp. 44-45). 

Differences from the superficially similar Hartlaubella gelatinosa are given under that 
species, and from the hydroid stages of other Obelia species by Cornelius (\915a : table 1). 

Mayer (1910), Stechow (1925) and recently Stepanyants (1979) regarded Obelia austro- 
georgiae Jaderholm, 1904#, as conspecific. I do not disagree (cf. Cornelius, 1975a) but have 
no fresh evidence. However, I cannot yet agree with Stepanyants that Clytia paulensis 
Vanhoffen, 1910, is identical with O. bidentata. The young medusae of both are sufficiently 
well described to indicate that two species are involved (see notes on Dispersive stage under 
the two species, pp. 89, 113). It could be argued that since Vanhoffen's material was 
infertile it was merely young O. bidentata: but this was not Stepanyants' argument. I believe 
neither Stepanyants nor I have seen the type material of C. paulensis. If it did prove to be 
young O. bidentata then a new name would have to be found for C. paulensis auct. (see 
synonymy in this paper). In that event it would be useful to apeal to the International 
Commission on Zoological Nomenclature to validate the widely used name paulensis sens, 

Clytia arborescens Pictet, 1 893 : 34-35, pi. 2, figs 30-3 1 , originally based on material from 
Indonesia, was reported south of Madeira at 100 m by Billard (1907); but Stechow (1925) 
included no further records in his synoptic list of warm water W Atlantic hydroids. Billard's 
material lacked gonothecae but, like the type material, had a compound stem and long 
hydrotheca (560-700 /zm) with sharply pointed cusps. It is likely that his material was O. 
bidentata; and it is relevant that he had wrongly identified O. bidentata from the English 
Channel (see Distribution, above). O. bidentata was not well known at the time he published 
the arborescens record. The type material of arborescens was probably not O. bidentata, 
however, since Pictet reported hemispherical medusa buds in the gonotheca, indicating 
Clytia sp. 

Obelia dichotoma (Linnaeus, 1758) 
(For illustrations see Cornelius, 1975a) 

Sertularia dichotoma Linnaeus, 1758 : 812. 

ICymodocea simplex Lamouroux, 1816 : 2 1 6, pi. 7, fig. 2 (see p. 121). 

ITubularia clytioides Lamouroux, in Freycinet, 1824 : 620, pi. 95, figs 6-8. 

Lomedea gracilis C. Pickering, in Dana, 1846 : 689, -pi. 61, figs 7, 7a-b (lapsus pro Laomedea; see p. 


Obelia dichotoma: Hincks, 1868: 156-157, pi. 28; Cornelius, 1975a : 265-272, figs 3-4. 
Obelia plicata Hincks, 1868 : 159, pi. 30, figs 1, la. 
Campanularia cheloniae Allman, 1 888 : 22, pi. 11, figs 2, 2a. 
Obelia dischotoma Billard, 1927 : 332 (lapsus pro dichotoma). 

lObelia commensuralis: Gudger, 1937: 1-6 (?lapsus pro 0. commissuralisMcCrady, 1857). 
Obelia hyaliana Vannucci, 1955 : 56. 
Laomedea plicata: Rasmussen, 1973 : 30, fig. 9. 

Eulaomedajlexuosa: Rees & Thursfield, 1965 : 102-103 (lapsus pro Eulaomedea\ redet.; see p. 105). 
For further synonymy see Remarks, Table 4 and Cornelius ( 1 975a). 


DISPERSIVE STAGE. Medusae of O. dichotoma and O. geniculata reared from the hydroids in 
the laboratory still cannot be distinguished and some taxonomic confusion continues 
(Russell, 1953; Cornelius, 1975a; Arai & Brinckmann-Voss, 1980; see also Remarks). 
Russell's book includes the best account of the biology of Obelia spp. medusae. 

The alternation of generations characteristic of medusoid coelenterates was actually first 
appreciated in scyphozoans, in 1829. It was reported in Obelia as early as 1836, first among 
the hydrozoans; but several still earlier workers had come close to its discovery in this genus. 
This and other historical aspects have been reviewed recently (Winsor, 1976; Cornelius, 
\911a, b). A paper by Desor (1849) was missed by these reviewers. Desor provided an early 
confirmation of medusa release in Obelia, probably O. dichotoma. However, like several 
contemporary workers (see p. 97) he assigned his material to Hartlaubella gelatinosa 
which has no medusa. He probably identified his material using the well illustrated work of 
van Beneden (1844), who made the same error. Several other workers have assigned an 
Obelia-\ike medusa to H. gelatinosa and, despite the early clarification by Couch (1844), 
there was confusion for many years between O. dichotoma, O. geniculata, H. gelatinosa and 
Gonothyraea loveni (see notes under H. gelatinosa (p. 97), and Cornelius, 1977a). 

Couch (1844) was in addition the first to note medusa release in Obelia geniculata (p. 120). 
Other early records in that species were contributed by F. W. L. Thomas (in Johnston, 
1847: 467), Hincks (1852 : 85)andGosse(1853 : 84-90, pi. 4). 

Illustrations of the medusae of 0. dichotoma were first provided by Baster (1762, pi. 5, fig. 
7), but the earliest of 0. geniculata appeared nearly a century later, in Gosse's (1853) book. 

Naumov (1969, as O. longissima, here referred to O. dichotoma) reported that 'the 
medusae were apparently not liberated from' Black Sea populations. But the species of 
Obelia are exceptionally well documented and since there is no other report of medusa 
retention in the genus a repeat of the observations would be desirable. 

REPRODUCTIVE SEASON. Medusae of Obelia spp. have been reported around the British Isles 
in all months but are commonest from 'spring to late autumn' (Russell, 1953). Fertile 
colonies were recorded on the coast of Egypt between 6 Sep and 4 Oct, 1933 (Billard, 1936). 

DISTRIBUTION. Found throughout European and African coastal waters with the exception 
of the Black Sea, from which there are no confirmed records (but see last paragraph of 
Dispersive stage). The species is remarkable in being nearly cosmopolitan in coastal waters 
and is one of the most widely distributed of hydroids. Aspects of this are discussed on page 
45; and a detailed summary of locality records has appeared elsewhere (Cornelius, 1975a). 

HABITAT. Tolerance of 12%o salinity recorded by Calder (1976) in S Carolina. Unusual 
substrates noted in the literature included a sponge and the fins of a shark (Couch, 
1 844 : XV, as 'the sea thread of Ellis'), a turtle (Allman, 1 888, as Campanularia cheloniae), a 
crab (Rasmussen, 1973: the colony was thought to be feeding on debris in the exhalent 
current); on the parasitic copepod Lernaeocera on the gills of a cod (Gadus); and on the 
back and claws of an anomuran 'crab', Lithodes maia (in Evans, 1978). Probably the record 
by Gudger (1937), of 'Obelia commensuralis' on the blueback herring (Alosa aestivalis), 
referred to this species; but his account suggested that the hydrorhizae entered the muscles of 
the fish and the identification is not certain. 

REMARKS. A revision of this species has appeared recently (Cornelius, 1975a) and only 
additional notes are given here. Identification of the hydroid stage is discussed above (p. 
1 13). Nematocysts were described by Ostman (1979) and by others whom she cited. 

Tubularia clytioides Lamouroux, in Freycinet, 1824, was based on live material collected 
from algae in the Azores. Probably it was Obelia dichotoma. Rees & White (1966) evidently 
thought Silicularia gracilis Meyen, 1834, also described from Azores material, to be 
conspecific but while the clytioides type specimen was an erect colony that of gracilis was 
reptant. Rees & White presented a muddled synonymy, referring to the 'species' 
(clytioides + gracilis) as Orthopyxis clytioides (Lamouroux); and including in its synonymy 


the entry "Silicularia clytioides Meyen, 1834', evidently a lapsus for "S. gracilis Meyen'. 
Nutting (1915) had made the same error fifty years earlier (p. 50). 

The once widely recognized nominal species O. commissuralis McCrady, 1857, based on 
the hydroid stage, was referred to O. dichotoma first by Bohm (1878), and again by me 
(19750). Bohm's paper has been overlooked; and I know of no other works giving this 

Obelia plicata Hincks, 1868, described from infertile hydroid material, type locality 
'Shetland', was recently referred to the present species without comment (Cornelius, 1975a). 
I have not been able to locate type material. The species deserves close evaluation as it has 
been recognized by several authors (Marktanner-Turneretscher, 1890; Nutting, 1915, as 
?O. plicata; Billard, 1927; Kramp, 1935, as Laomedea plicata; Fraser, 1944; Teissier, 1965; 
Rasmussen, 1973) although not by Broch (19 126). O. plicata was said by Hincks to resemble 
H. gelatinosa in having compound stems but the hydrothecal rims were even, not castellated. 
The hydrotheca was said by Hincks 'very much [to] resemble in form that of O. dichotoma". 
Gonothecae were first assigned to O. plicata by Marktanner-Turneretscher (1890) who said 
they were the same as those of O. dichotoma. Indeed, the two species seem conspecific. Some 
older colonies of O. dichotoma (e.g. BMNH 1948.9.8.14, Plymouth, 21 Apr 1899, coll. E. T. 
Browne) exhibit compound stems resulting from overgrowth of up to 10 young, pale stems 
along the older, thicker and darker ones. The specimen mentioned resembles closely the 
colony illustrated by Hincks as O. plicta, and synonymy seems justified. Godeaux (1941, as 
H. gelatinosa) described similar O. dichotoma material, from the North Sea; while Billard 
(1927) maintained O. plicata distinct on the basis of the original characters. Broch (19126) 
alone raised doubts, but was non-committal. 

The factors causing overgrowth in occasional O. dichotoma specimens are not known, 
most colonies being solitary. However, such overgrowth has been described in other hydroids 
and called 'auto-epizoism' (Millard, 1973). If an overgrown specimen results from second 
and subsequent planulae settling on the original colony then the whole should be regarded as 
an aggregation of colonies, and not as a single colony. The phenomenon is discussed above 
(p. 40). 

Campanularia cheloniae Allman, 1888, was based on infertile material collected during 
the 'Challenged expedition from the back of a turtle; but the locality of the turtle was not 
recorded. The original description and what remains of the type series (BMNH 
1 888. 1 1.13.15, small colony in spirit + microslide preparation) are probably O. dichotoma. 

Laomedea sargassi Broch, 1913, was recorded from Ghana by Buchanan (1957 : 360). 
The name sargassi was a nom. nov. for O. hyalina Clarke, 1879, introduced just to prevent 
confusion with Gonothyraea hyalina auct., and is therefore inadmissible under the current 
conventions of nomenclature. Congdon (1907 : 468) described material from Bermuda as O. 
hyalina, but Hargitt (1909) gave that material specific status, as Laomedea congdoni. I have 
previously (Cornelius, 1975a) referred O. hyalina Clarke and O. congdoni (Hargitt) to 
O. dichotoma, but overlooked the nom. nov. sargassi. Recent examination of the type 
material of O. congdoni (U.S. nat. mus. cat. no. 42647, microslide preparation of infertile 
fragments) confirms that it too is simply O. dichotoma. Obelia hyaliana Vannucci, ? in ms, 
was a name given to some "O. hyalina' material which Vannucci (1955) later considered 

Obelia geniculata (Linnaeus, 1758) 
(For illustrations see Cornelius, 19750) 

Sertularia geniculata Linnaeus, 1758 : 812. 

Obelia geniculata: Hincks, 1868: 149-151, pi. 25, fig. 1, la; Cornelius, 1 9750:272-278, figs 1,5. 

For further synonymy see Cornelius ( 1 91 5a), Remarks and Table 4. 

DISPERSIVE STAGE. A medusa. See Remarks and notes under Obelia dichotoma. 

REPRODUCTIVE SEASON. Russell (1953) recorded the following in British waters: Obelia sp. 
medusae found nearly throughout year, especially 'spring to late autumn' (Russell); medusae 


released at Plymouth, March-November (Orton, 1920); some evidence of lunar periodicity 
in release times (Elmhirst, 1925). 

In the Mediterranean, Lo Bianco (1909) recorded medusa release at Naples March-June 
and October-January. 

DISTRIBUTION. An almost cosmopolitan shallow water species found nearly throughout the 
eastern North Altantic. 

The species was not recorded from the Black Sea by Naumov (1969). He listed the species 
as absent from the Baltic as well but Stechow (1927) reported it from Stockholm. It was 
recorded from Iceland and the Faeroes but not from Spitzbergen by Kramp (1929), although 
already known from Jan Meyen Island (Broch, 1918). Calder (1970) among others recorded 
the species from W Greenland; but I (Cornelius, 19750) misquoted him as including E 
Greenland in its distribution. I know of no records from E Greenland. 

HABITAT [see also Geographical distribution (p. 45), the habitat notes under O. dichotoma 
and Cornelius, 19750]. The species was collected from the fins of a spur-dog (Squalus 
acanthias) by Couch (1844), who deduced that planula settlement must be rapid. Other 
actively swimming animals on which O. geniculata has been recorded include a sea-horse 
(Hippocampus guttulatus, by Zirpolo, 1939, 1940) and parasitic copepods themselves on 
fish (Lernaeocera spp. on Gadus (cod), by Leloup, 193 1/?; on Lernaeocem on Clupea 
(herring), by Debouteville & Nunes, 1951, 1952; on Dinemoura on Cetorhinus (basking 
shark) by Debouteville & Nunes). Long distance travel on floating algae has also been 
recorded (Arnaud, Arnaud, Intes & Le Loeuff, 1976). Recorded at 100 m depth by Naumov 
(1960). However, some of these records on fish might have been of 0. dichotoma. 

REMARKS. A revision of this species has appeared elsewhere (Cornelius, 19750) and the 
present account simply adds to that. 

Couch (1844) gave some interesting details. He stated the number of tentacles of the 
hydranth to vary widely (1 1-29 on p. 2, 14-28 on p. 38). He described medusa bud develop- 
ment but not medusa release, which was not known till 1847 in this species (Cornelius, 
19750, 1977a, b; see also p. 1 18). He recorded a range of tissue colour in the hydroid from 
'white to deep red'. 

The nematocysts were described by Ostman (1979) and by others whom she cited. 

Campanularia vermicularis Van Beneden, 1866, was described without illustrations and 
identification has been thought impossible (Vervoort, 19460; Leloup, 1947). The species was 
based on Belgian material. Van Beneden kept live colonies for three weeks. He described the 
species as being more robust than O. geniculata, and as having larger hydrothecae and more 
numerous annulations. It was collected off the brown alga, Fucus vesiculosus L. The 
hydranth was said to have as many as 40^4 tentacles, a character which might one day help 
to identify his material (cf. Couch's observations, immediately above). Meanwhile I follow 
Vervoort in treating the species here but not including it in the synonymy. 

Obelia spp. indet. 

1. Hydroids. Obelia longicyatha Allman (1877 : 10, pi. 7, figs 4-5) was recorded from 
35 42' N, 8 40' W (Cap Spartel, Tangier) and the Gulf of Cadiz by Billard (1907 : 168, as 
Clytia longicyatha), but the identity of the species remains unclear. I have discussed it 
elsewhere (Cornelius, 19750:264). See also the note on Campanularia vermicularis Van 
Beneden, 1866 (immediately above). 

2. Medusae. Some species from the eastern North Atlantic cannot be assigned to a hydroid. 
Obelia adriatica Neppi (1912 : 726-727, pi. 3, fig. 8, 8a) was described from Adriatic 
material; and Obeliopsis fabredomergui Le Danois (1913:110) was based on material from 
the Little Minch, NW Scotland. It seems most unlikely that either is valid. O. plana Sars 
(1835 : 28, pi. 5, fig. 13, as Thaumantias) was referred to O. jlabellata hydroid by Bedot 
(1910 : 484) and some later authors, and hence would fall in the synonymy of O. dichotoma. 


But there seems slim reason for this identification (cf. Cornelius, 19750:271, footnote). 
Other pre-1910 medusa names in this genus were listed by Bedot (1901-1925) and Mayer 
(1910), and are not repeated here. As explained above (p. 1 18) most cannot be confidently 
assigned to a hydroid stage and a new list would serve little purpose. 

Taxa now referred to other families 

A few genera and species recorded from the eastern North Atlantic are no longer referred to 
the Campanulariidae. As several have been included in this family in some standard works 
notes on them are given here to summarize the current opinions. The taxa are treated in date 

1. The genus Capsularia Cuvier, 1797, was used by Gray (1848:85-87) to embrace 
several British species of Campanulariidae but should be regarded a junior synonym of 
Coryne Gaertner, in Pallas, 1774 (discussion in Cornelius, 1 9756 : 378). 

2. Sert ularia fruticosa Esper, 71810 [pi. 34, figs 1-2 (see note 2, p. 124); also Hammer, in 
Esper, 1829:162-163 (syn. Sertularia laxa Lamarck, 1816:116; Laomedea sauvagii 
Lamouroux, 1816: 206; both nom. nov. pro S. fruticosa)] was recorded from the Adriatic Sea 
by Marktanner-Turneretscher (1890 : 205) as Campanularia fruticosa. The species is now 
referred to the genus Thyroscyphus in the family Sertulariidae (e.g. Vervoort, 1967; Millard, 
1975). It is otherwise unrecorded from the eastern North Atlantic and Marktanner- 
Turneretscher's record may well be wrong. However, T.fruticosus occurs in the Red Sea as 
far north as the Gulf of Suez (Vervoort, 1 967). 

The exact date of publication of the species name is unusually difficult to determine and 
the necessary bibliographic work on Esper's important multi-part book has apparently not 
been done. Precise dating would be useful as the plates were issued several at a time around 
1810, and they carry binominals. They thus predate the posthumous text, edited by Hammer 
(Esper, 1 829), from which this part of the work is often thought to date. The plates predate 
also several important European works on zoophytes [e.g. those by Lamarck (1816) and 
Lamouroux (1812, 1816, 1821; Lamouroux et al., 1824)]. Esper's work is rare, and few of 
the original wrappers survive; but the information on an incomplete set of wrappers in the 
BMNH library gives hope that dating would be simple if a complete copy were located. See 
also section 7, below. 

3. The genus Cymodocea Lamouroux, 1816:214, originally included two species, C. 
simplex and C. ramosa, both described as new. C. simplex was based on material from Great 
Yarmouth and 'Ireland', C. ramosa on fragments from the Antilles. Johnston (1838) 
commented on the genus and on the identities of the two species; and Gray (1848) dismissed 
them as 'doubtful species' at the end of the Campanulariidae. Lamouroux (1821) and 
Lamouroux et al. (1824) included additional species in the genus but these do not enter the 
present discussion. Billard (1909), who saw some of Lamouroux' original material, referred 
the two species to Nemertesia antennina (Linnaeus, 1758), family Plumulariidae. I 
designate C. ramosa type species of the genus Cymodocea, the name of which hence 
becomes a junior subjective synonym of Nemertesia Lamouroux, 1812. Billard was confi- 
dent that the C. ramosa material he saw was type, but had doubts about that of C. simplex. 
His doubts are significant since in J. Fleming's opinion, quoted by Johnston (1838), the 
original illustrations of C. simplex might be identified as an Obelia dichotoma colony lacking 
hydrothecae. I concur with Fleming's opinion; and disagree with Johnston who thought 
simplex was a plumularid. If the illustration is actually of O. dichotoma then Billard's 
caution was justified and the specimen he saw was not type. Almost all the Lamouroux 
collections were destroyed during the liberation of Caen on 7 July, 1944, so the C. simplex 
material is no longer available (Redier, 1967). However, C. simplex is here provisionally 
referred to O. dichotoma (p. 1 1 7). 


The species Cymodocea comata Lamouroux, 1821, was recorded from the coast of Devon 
by W. E. Leach (in Johnston, 1838); but the species was referred to Nemertesia ramosa 
(Lamarck, 1 8 1 6) by Billard ( 1 909), who saw type material. 

4. Campanularia intertexta Couch, 1844 : 41^2, pi. 11, fig. 3, was based on material 
then in the Museum of the Royal Institution of Cornwall, Truro. The material almost 
certainly no longer exists (Curator, Roy. Inst. Cornwall, pers. comm.). Johnston 
(1847:109-110) repeated Couch's description verbatim but regarded the species as 
unsoundly based. Gray (1848 : 88) listed it without comment, incidentally assigning to it 
Shetland material of Lafoea dumosa (Fleming, 1820), BMNH reg. no. 1847.9.24.66. At 
the same time Gray proposed the new genus Conchella (?lapsus pro Couchella), of which 
Campanularia intertexta is type species by monotypy. Hincks (1868:220) thought the 
species might be L. dumosa, stating that Couch's description was inadequate for 
identification. Later Bedot (1905 : 157) referred intertexta to 'Coppinia arcta\ an invalid 
species accepted by several nineteenth century authors but merely based on the coppinia or 
reproductive branch of L. dumosa. Couch's illustration shows an unidentifiable campanu- 
larid hydroid, possibly Orthopyxis Integra, growing on a coppinia of L. dumosa. Thus the 
type 'series' was mixed. The name intertexta is here restricted solely to the illustrated 
coppinia, which becomes lectotype. Hence Campanularia intertexta is to be regarded a 
junior synoym of L. dumosa; and the genus name Conchella a junior synonym of Lafoea 
Lamouroux, 1821. See also page 65. 

5. Campanularia lacerata Johnston, 1847:111, pi. 28, fig. 3, is now known as 
Opercularella lacerata and referred to the Campanulinidae (e.g. Millard, 1975). The species 
has been referred to a variety of genera in the literature, including Capsularia, Laomedea, 
Wrightia and Calycella (references in Hincks, 1868). 

6. Laomedea obliqua Johnston, 1847 : 106-107, pi. 28, fig. 1, based on British material, is 
today known as Monotheca obliqua and Plumularia obliqua by different authors and 
referred to the Plumulariidae (e.g. Millard, 1975). 

7. Campanularia fruticosa Sars, 1850 : 138-139, is today referred to Lafoea dumosa 
(Fleming, 1820), in the Lafoeidae (e.g. Cornelius, 19756). It has often been given full 
specific status, as L. fruticosa. See also section 2, above. 

8. Campanularia abietina Sars, 1850 : 139, based on Norwegian material, has long been 
known as Grammaria abietina and assigned to the family Lafoeidae (e.g. Cornelius, 19756). 

9. Campanularia parvula Hincks, 1853 : 178, pi. 5a, was probably based on material of 
Calycella syringa (Linnaeus, 1767) lacking operculae. Several authors have referred the 
species to Lafoea Lamouroux, 1821, but this seems wrong (references and discussion in 
Cornelius, 19756). 

10. Campanularia gracillima Alder, 1856a:361, pi. 14, figs 5-6, based on hydroid 
material from NE England, has frequently been referred to Lafoea, family Lafoeidae; but 
lately to the species Lafoea dumosa (Fleming, 1820) (e.g. Cornelius, 19756; Cornelius & 
Garfath, 1980). 

11. Laomedea acuminata Alder, 18566 : 441, pi. 16, figs 5-8, based on hydroid material 
from NE England, is the hydroid of an Aequorea sp. medusa, family Aequoreidae. The two 
species recognized from British waters from the medusa stage were described earlier so the 
name acuminata should be regarded provisional. Although the medusa generation of the two 
can be separated the respective hydroids are morphologically identical, so far as is known. 
Hence at present it cannot be decided to which of the 'medusa species' acuminata should 
rightly be referred (Russell, 1953). 

12. The nominal species Laomedea tenuis Allman, 1859 : 367-368, was once known as 
Leptoscyphus tenuis (e.g. Hincks, 1868). It is currently regarded as a campanulinid, having 


been provisionally referred to Phialella quadrata (Forbes, 1848) by Stechow (1923a : 129) 
and Rees (1939:441). Browne (1896:479) summarized the mistaken observations by 
Allman who assigned the medusa of one species to the hydroid of another. Campanulina 
tennis Van Beneden, 1886 : 174, 176, pi. 13, is a different nominal species, provisionally 
referred to Aequorea vitrina Gosse, 1853, by Russell (1953). Bedot (1910) listed an earlier 
homonym of C. tenuis. 

13. Campanularia fastigiata Alder, 1860 : 73-74, pi. 5, fig. 1, based on Scottish material, 
was known for some decades as Stegopoma fastigiatum. Edwards (1973) showed that 
fastigiatum was the hydroid stage of the medusa Modeeria rotunda (Quoy & Gaimard, 

1827), the binominal of which takes priority. The species is assigned to the family 
Laodiceidae (e.g. Rees & Rowe, 1969; Edwards, 1973; see also Cornelius & Garfath, 1980). 

14. Campanularia humilis Hincks, 1866 : 298, is now known as Cuspidella humilis. Its 
affinities are unclear but it was removed from the Campanulariidae long ago (discussion in 
Cornelius & Garfath, 1 980). 

15. Clytia eucophora Haeckel, 1879 : 168, was a combination applied to the hydroid stage 
of Eucopium primordiale Haeckel, 1879 : 168 (sic), from Corsica. As Mayer (1910 : 236) 
pointed out, the two names are objective synonyms with the same date of publication. 
Mayer, as first reviser, suppressed eucophora in favour of primodiale\ but at the same time 
referred Haeckel's species to Eucope picta Keferstein & Ehlers, 1861. Although Kramp 
(1961) omitted to treat E. picta there is no doubt that it and the Haeckel species should be 
referred to the Phialellidae. Weismann (1883 : 158) introduced the combination Clytia 
eucopophora, a lapsus of Haeckel's spelling. 

16. Campanularia mutabilis Ritchie, 1907 : 504, pi. 23, figs 3-5, based on Azores 
material, is now known as Scandia mutabilis and referred to the Lafoeidae (e.g. Millard, 

17. Campanularia divisa was attributed by Bassindale (1941 : 148) to Todd (1906 : 137) 
whom he supposed to have reported material from Ilfracombe. Bassindale misread Todd's 
list, which included Campanularia species and Tubularia indivisa. The word 'indivisa' was 
split between the two lines (in/divisa) and evidently read wrongly by Bassindale. 

18. The genus Hincksella Billard, 1918 : 22, was considered by Totton (1930) and Ralph 
(1957) to be closely related to the Campanulariidae, but is now referred to the Syntheciidae 
(e.g. by Millard, 1975). 

19. The genus Billardia Totton, 1930 : 150 (type species B. novae zealandiae, by original 
designation) was based on southern hemisphere material. Totton assigned Billardia to the 
Campanulariidae but the genus is similar to Hincksella and like it comes within the scope of 
the Syntheciidae sensu Millard (1975), in my opinion. Blanco (\961b) and Stepanyants 
(1979), however, retained Billardia in the Campanulariidae. Possibly a greater under- 
standing of the reproductive structures will help. Totton suspected that the blastostyles of 
Billardia were produced in place of hydranths, within the hydrothecae, but said his material 
was inadequately preserved for him to be certain. If his suspicion were confirmed the 
Syntheciidae would be the correct family for Billardia. 


1 (See p. 65). G. D. Westendorp (1813-1868) apparently produced only two works 
involving coelenterates (Westendorp, 1843, 1853). Both were on the zoophytes of the 
Belgian coast. The first was a straightforward taxonomic account including some new 
genera and species, among them Clytia ryckholtii (here referred to Orthopyxis integrd). 
Probably none is valid. His second work, published in 1853, was remarkable in being 
illustrated by dried specimens mounted on sheets. It is rare but still important since one of 


the included species was described as new in the 1843 work, and the specimens used in 
illustration might be considered types. The 1853 work comprised a printed title page, a 
printed page giving a list of species and localities, and 32 hebarium sheets each bearing one 
species. There were 16 hydroids, 14 bryozoans and 2 sponges. Examples of the work were 
seen by Neviani (1903), Bedot (1910 : 200-201) and Billard (1914; also seen by Leloup, 
1947 : 5); and another, imperfect copy has recently been found in the British Museum 
(Natural History) (Cornelius, unpublished ms in BMNH). I have not seen an intact copy. 

Neviani listed the 32 species but C. ryckholtii was not among them. The copy described by 
Billard (1914), in Brussels (Mus. roy. Hist, nat.; inv. no. 3440), similarly does not have C. 
ryckholtii. The BMNH example also does not include that species. The copy evidently 
came to the (then) British Museum library about 1867. Certainly a collection of specimens 
corresponding with Neviani's list was purchased from Westendorp at that time, and was 
given the 32 zoological accession numbers 1867.5.4.22-24 and 1867.5.7.1-29. The 
specimens were curated each under its own species, and the printed species list was cut up to 
provide labels which were in most cases glued to the herbarium sheets. Much, perhaps all, of 
this material survives and is still curated under the various species; but I have not found the 
title page of this copy. 

Only one type specimen of any group is included in this collection, that of the hydroid, 
Plumularia macleodi Westendorp, 1843 (BMNH reg. no. 1867.5.7.7). It can be considered a 
syntype of P. macleodi; and is in fact a specimen of Kirchenpaueria pinnata (Linnaeus, 
1758). P. macleodi is thus a junior synonym of K. pinnata. Billard (1914), working on the 
corresponding Brussels syntype specimen, reached the same conclusion. 

2 (See p. 121). Some evidence was presented by Sherborn (1922 : 1, entry under Esper; 
Sherborn, 1926 : 2528, entries under fruticosa, Laomedea and fruticosa, Sertularia) that at 
least some of the plates of Esper's (1829) 'Die Pflanzen-Thiere' appeared before 1816, 
possibly around 1810. Since the plates bear binominals, and might predate the synoptic 
works of Lamouroux (1812 onwards) and Lamarck (1816 onwards), correct dating is 
important. However, it seems that bibliographers have not yet dated the plates and 
Sherborn's (1926) provisional date of 1810 for the plate of Sertularia fruticosa Esper is the 
best compromise. Certainly the plate of fruticosa predated the works of Lamarck ( 1 8 1 6) and 
Lamouroux (1816), which referred to Esper's plate as being already published. Indeed, they 
proposed new names in place of fruticosa. The three volume Esper work was issued in parts, 
some after Esper's death in 1810; and the few copies I have seen have been collated with the 
undoubtedly earlier Sertularia plates intermingled with the text. But the partial set of 
wrappers with the BMNH copy gives much detailed information, and a full set might 
enable all the plates of this scarce work to be dated accurately. 

3 (See p. 1 12). Dating of the two works here listed as Peron & Lesueur, 1810a, 718106, has 
caused confusion. A clarification is desirable since in them many genus names of medusae 
were introduced which are still used. The works comprised the earliest serious systematic 
treatment of medusae, and these authors finally disbanded the genus Medusa Linnaeus, 
1758. The first of the two works was one of a series of journal papers written by the two 
authors, and comprised their taxonomic treatment of medusae. Several other papers in the 
series also dealt with marine animals but are not important here. They have been listed by 
Goy (1980). The second work (Peron & Lesueur, 718106), a book entitled Histoire generale 
des meduses, brought the series of papers together each forming a chapter of the book. The 
book version was repaginated from 1 on, and was presumably published after the series of 
papers (although it might conceivably have been issued in parts as the papers appeared). 

The publication date of the taxonomic paper (Peron & Lesueur, 1810#) has been taken as 

1809 by most authors and this date might be inferred from the title page of the volume of the 
journal in which it appeared. But there is good evidence that it did not appear until January, 

1810 (Sherborn, 1914; 1929 : 4455, entry under Obelia\ Cornelius, 1975a; Goy, 1980). The 
book, the Histoire generale des meduses, has been almost entirely overlooked. It was dated 


1809 in the British museum catalogue of printed books. But J. Goy (pers. comm.) and I 
concur that the book version almost certainly would have followed the series of papers, and 
so would have appeared at least as late as 1810 (unless the book version were issued in parts). 

Hence the many names introduced by Peron & Lesueur almost certainly date from the 
1810^ paper, published January, 1810. Apart from minor heading changes the book version 
(71810&) was apparently printed from the same 'blocks' as the series of papers. For 
taxonomic purposes the two versions differ only in date. 

The numerous plates that were to have accompanied the account of the medusae (Peron & 
Lesueur, 1810#) were prepared but not published immediately (cf. Cornelius, \915a : 253, 
footnote; 1977a:49, footnote). But most of the illustrations showing Peron & Lesueur's 
'new' species were brought together in an extremely rare work usually attributed to Lesueur 
alone (Peron & Lesueur, 71815). This work was cited by Haeckel (1879, in several of his 
synonymies, as the Planches inedites) and Totton (1965 : p. 45, pi. 8) among others but 
almost all twentieth century medusa workers have overlooked it. Fewer than half a dozen 
copies are known (Totton, 1965; Goy, 1980). Goy (1980) cited evidence from the P.-v. 
Seanc. Acad. Sci. Paris, 1795-1831 (issue covering 14 August 1815) 5 : 532, that 1815 is the 
correct date; and not 71811 as given in the British museum catalogue of printed books 
(187 : 1 1 1) under Peron alone. She concluded that the bulk of the illustrations, showing most 
of the nominal species newly described by Peron & Lesueur (1810(2), were copied and 
published by a variety of contemporary compilers in their own works and so made public. 
They included such famous names as de Blainville, Cuvier, Milne Edwards, Lamarck and 
Lesson (references in Goy, 1980). The original Lesueur drawings are preserved in the 
Museum at Le Havre. 

Although the bulk of the rare work (Peron & Lesueur, 71815) comprised illustrations of 
medusae exquisitely drawn by Lesueur, the title page leaves no doubt that the authorship 
should be ascribed jointly to Peron & Lesueur (Mrs A. Datta, pers. comm.). Peron had died 
in 1810 and it can be inferred that Lesueur wished him still to be senior author, as he had 
been of the lengthy text (Peron & Lesueur, 1 8 1 Oa, 7 1 8 1 06) of the work. 


I am indebted to Professor W. Vervoort, Rijksmuseum van Natuurlijke Historic, Leiden, for 
generous and detailed discussion on the Campanulariidae; and through him to the Stichting 
Jan Joost ter Pelkwijk Fonds for financial assistance while on a visit both to the Leiden 
Museum and the Institut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit 
van Amsterdam, where Dr R. W. M. van Soest was my kind host. Dr J. M. Paul, Zoology 
Department, University of Oxford, helped trace literature about Laomedea neglecta. I am 
grateful also to Dr R. G. Hughes, Westfield College, University of London, for new records of 
Clytia paulensis and to Dr R. B. Williams of Tring for helpful discussions. Dr A. G. Long, 
Hancock Museum, Newcastle upon Tyne, kindly provided facilities while I worked on the 
Joshua Alder collection. Lastly I am grateful to Miss L. J. Wadmore for preparing the 
illustrations of medusae and to R. H. Harris for histological assitance. Other acknowledge- 
ments are made in the text. 


On 15 August 1981 I found Laomedea angulata (p. 98) common on the eel grass bed at 
Studland, Dorset. The previous record from the British mainland was dated 1906; and from 
Dorset was 1890, also at Studland. Whether or not the species had disappeared from that 
locality in the meantime might be hard to tell. On 15 September 1981 I found the same 
species abundant on eel grass near Misery Point, R Yealm, near Plymouth. At this locality 
too the status of the species during the last several decades was unclear. 



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Manuscript accepted for publication 24 July 1981. 


Valid names are in Roman type, new names in bold and invalid names in italics. New combinations are 
indicated by an asterisk (*). Main page references are in bold. References to substrates are not included. 

abietina, Campanularia & Grammaria 1 22 

acuminata, Campanulina 74 

acuminata, Laomedea 122 

acuta, Campanularia 74, 81 

adelungi, Obelia 1 1 4 

adriatica, Obelia 1 20 

Aequorea 122 

vitrina 123 
affinis, Clytia 7 1 
affinis, Eucope 11, 73, 74 
africana, Campanularia 60 
africana, Orthopyxis 39, 60 
agas, Campanularia 54 
Agastra 48, 57-58 

caliculata 6 1 

mira 6 1 , 67 

aha, Campanularia 53, 54 
alternata, Clytia 84, 85, 86 
alternata, Eucope & Obelia, 1 14 
andersoni, Obelia 1 14 
angulata, Campanularia & Eulaomedea 98 
angulata, Laomedea 41, 42, 43, 44, 98-102, 

104, 105; Fig. 17, p. 99 
angulosa, Obelia 1 14 
antennina, Nemertesia 121 
arborescens, Clytia 1 13, 1 14, 117 
arcta, Coppinia 122 
armata, Oceania 8 1 
arruensis, Obelia 1 14 
articulata, Eucope 1 14 
asymmetrica, Orthopyxis 6 1 , 67 
atlantica, Hypanthea 50 
atlantica, Silicularia 50 
attenuata, Campanularia 73-74, 79, 81 
attenuata, Clytia 40, 42, 8 1 
attenuata, Obelia 1 14 
australis, Obelia 1 14 
austrogeorgiae, Obelia 1 13, 117 

bicophora, Clytia (Trochopyxis) 73, 80 

bicophorum, Phialidium 74, 80 

bicuspidata, Obelia 90, 113, 1 14 

bidentata, Obelia 40, 43, 44, 85, 96, 97, 1 10, 

bifurca, Obelia 11 4, 115 
bifurcata, Obelia 1 14 
bilabiata, Campanularia 58, 59, 60 
bilabiata, Eucopella 58 
bilabiata, Silicularia 60 
Billardia 123 
novaezealandiae 123 

biserialis, Obelia 1 14 

bistriata, Laomedea 84, 85, 86 

borealis, Campanularia 6 1 , 66 

borealis, Obelia 1 14 

brachycaulis, Campanularia 53, 55-56, 74, 82 

brasiliensis, Campanularia 1 14 

braziliensis, Obelia 1 14 

breviscyphia, Campanularia 60-6 1 , 66 

brochi, Laomedea 47, 107, 110 

brochi, Paralaomedea 47 

buskiana, Thaumantias 74 

buskianum, Phialidium 74 

calceolifera, Campanularia 102 

calceolifera, Eulaomedea 98, 102 

calceolifera, Laomedea 41, 42, 43, 44, 47, 98, 

100, 101, 102-105; Fig. 18, p. 103 
calceolifera, f. of Laomedea angulata 100 
calceolifera, Obelia 102 
caliculata, Agastra 6 1 
caliculata, Campanularia 51, 58, 60-61, 64, 


caliculata, Eucopella 58,61 
caliculata, Orthopyxis 48, 6 1 , 63, 65-66 

syringa 55, 57, 78, 122 
calyculata, Campanularia 51,61 
Camp alar ia 97 

conferta 102-105 
campanella, Medusa 73 
Campanula 51 

Campanularia, Eucopella 58, 60, 61 , 63, 67 
Campanularia 4 1 , 47, 48, 49, 50, 51-52, 57, 70, 

abietina 122 

acuta 74, 81 

africana 60 

agas 54 

angulata 98 


bilabiata 58, 59, 60 

borealis 6 1 , 66 

brachycaulis 53, 55, 56, 74, 82 

brasiliensis 1 1 4 

breviscyphia 60, 6 1 , 66 

calceolifera 102 

caliculata 5 1 , 58, 60, 6 1 , 64, 65-^6 

calyculata 51,61 

caulini 1 14 

cavolinii 1 14 

cheloniae\\4, 117, 118,119 



compressa 60, 6 1 , 64, 66-67, 73 

confer ta 100, 104 

coruscans 1 14 

crenata 52-53, 59, 60; Fig. 2, p. 55 

crenata f. intermedia 59 

decipiens 107 

devisa 123 

edwardsill, 81 

elongata 107,110 


fastigiata 123 



flexuosa 105 

fragilis 107, 109,110 

fruticosa 121, 122 


gelatinosa 95 

geniculata sens. Lister 9 1 

gigantea 73, 75,81 

gracilis sens. Allman 60, 61, 66; sens. Meyen 

50;sens. Sars 74, 77,81 
gracillima 122 
groenlandica 52, 55, 56, 57 
hicksoni 54 

hincksii 5 1 , 53-55, 80, 82; Fig. 3, p. 55 
humilis 123 
hummelincki 82 
inconspicua 73 
Integra 52,60 
integriformis 6 1 , 64, 66-67 
intermedia 58,60 
inter text a 65 , 122 
johnstoni 71 , 73, 75, 77, 79-80 
kincaidi 9 1 
lacerata 122 
lennoxensis 58, 60 
macrotheca 53,54 
magnifica 52,53 
maior 1 1 4 
morgansi 5 1 
mutabilis 123 
neglecta 107 
noliformis 80 
obliqua 84 
obtusidens 1 14 
parvula 122 
paulensis 88 
pelagica 73, 78-79 
platycarpa 6 1 
prolifera 1 14 
ptychocyathus 80 
pulcratheca 5 1 
rara 53 

raridentata 73, 74, 75, 76, 80, 8 1 , 90 
retroflexa 5 1 

speciosa 52, 53 
speciosus 52 
spinulosa 1 14 

tulipifera 70 

tulpifera 70 

vermicularis 120 

verticellata 67 


villafrancensis 74, 81 

volubiliformis 73, 80 

volubilis 48, 5 1 , 54, 55-57, 69, 70, 73, 76, 77, 

79; Fig. 4, p. 55 

CAMPANULARIINAE 48-49, 50-51, 69-70 
campanulata, Eucope 7 1 , 73, 74 
Campanulata 5 1 

verticillata 68 
Campanulina acuminata 74 

tennis 123 
Capsularia 121, 122 

Integra 6 1 

/aev/5 6 1 , 65 
carnea, Podocoryne 43 
caulini, Campanularia 1 14 
cavolinii, Campanularia 1 14 
cheloniae, Campanularia 1 14, 1 17, 1 18, 1 19 
chinensis, Obelia 1 14 
Clytea 70 

vicophora 73 
Clythia 57, 70 
C/y//z/ajohnstoni 74 

poterium 6 1 , 66 
Clytia 41 , 42, 47, 48, 49, 57, 69, 70-72 

affinis 1 \ 

alter nata 84,85,86 

arborescens 1 13, 1 14, 117 

attenuata 40,42,81 

bicophora 73,80 

compressa 14, 82 

cylindrica 42, 73, 80 

discoida* 42, 72-73; Fig. 8, p. 72 

edwardsi 8 1 

ed wards ia 43 

elsaeoswaldae 80 

eucophora, eucopophora 123 

flavidula 73, 74, 76, 77, 80, 81 

folleata 9 1 

foxi 84, 86 

gardineri 9 1 

geniculata 84, 86 

gigantea 40, 74, 77 

gracilis 42, 71,91 


hemisphaerica 40, 41, 42, 43, 45, 70-71, 
73-82, 86, 87, 89, 90, 91, 92, 94; Fig. 9, 
p. 75 

hendersonae 84 


hummelincki 4 1 , 42, 44, 69, 82-83, 86; 
Fig. 10, p. 82 

islandica* 42, 84; Fig. 11, p. 84 

johnstoni 69, 70, 73, 74, 80 

kincaidi 9 1 

teem 55, 57, 73 

tanguida 8 1 

linearis* 42, 84-86; Fig. 12, p. 85 

longicyatha 120 

longiiheca 1 1 4 

mccradyi* 42, 87-88; Fig. 13, p. 87 


o///brw/.s73,80, 81 

obeliformis 13,15, &\ 


olivacea 67 

Clytia (Orthopyxis) poten'MW 57, 58,60,61,66 
Clytia paulensis 42, 43, 44, 76, 85, 86, 88-91, 
113, 115, 117; Fig. 14, p. 89 

pelagica 73 


Clytia (Platypyxis) 70, 71, 80 
Clytia posterior 6 1 


rijckholtii 6 1 , 65 

ryckholtii 6 1,65, 123-124 


serrata 85 

serrulata 73, 74, 76 

simplex 1?>,&\ 

striata 86 

Clytia (Trochopvxis) 70 
Clytia ulvae 88, 91 

undulata 60, 6 1 , 65 

unijlora 74 

urnigera 64-65, 73, 78 

vicophora 73 

villafrancensis 74 

viridicans 74 


volubis 43 

CLYTIINAE 38, 48-49, 50, 69-70 

clytioides, Orthopyxis 50, 118-119 
clytioides, Silicularia 50, 1 18-1 19 
clytioides, Tubularia 50, 1 14, 1 17, 118 
Cmpanularia 5 1 

comata, Cymodocea & Nemertesia 121, 122 
commensuralis, Obelia 1 14, 1 17, 118 
commissuralis, Obelia 1 14, 1 17, 119 
compressa, Campanularia 60, 6 1 , 64, 66-67, 73 
compressa, Clytia 74, 82 
compressa, Orthopyxis 6 1 , 66 
ConcHella 122 

intertexta 122 

conferta, Campalaria 102-103, 104-105 
conferta, Campanularia 100, 104 
conferta, Laomedea41, 102-103, 104 
congdoni, Laomedea& Obelia 1 14, 119 


convexa, Thaumantias 73 

Coppinia arcta \ 22 

corona, Obelia 1 14 

coruscans, Campanularia 1 14 

Coryne65, 122 


coughtreyi, Obelia 1 14 

crenata, Campanularia 52-53, 59, 60; Fig. 2, 

p. 55 

crenata, Eucopella 52, 58 
crenata, Orthopyxis 40, 42, 52, 58-60; Fig. 5, 

p. 59 

Cuspidella humilis 123 
cylindrica, Clytia 42, 73, 80 
cylindrica, Platypyxis 73 
cymbaloidea, Epenthesis 73, 74 
cymbaloidea. Medusa 73 
Cymodocea 1 2 1 

comata 122 

ramosa 1 2 1 

simplex 114, 117,121 

decipiens, Campanularia 107 
decipiens, Laomedea 107, 109, 110 
delicata, Orthopyxis 58, 59, 60 
diaphana, Eucope & Thaumantias 1 14 
dichotoma, Obelia 40, 41, 43, 45, 73, 96, 100, 

101, 102, 104, 112, 113, 114, 117-119, 


dichotoma, Sertularia \ 1 7 
dischotoma, Obelia 1 14, 1 17 
discoida,* Clytia 42, 72-73; Fig. 8, p. 72 
discoida, Oceania 72 
discoidum, Phialidium 73 
divaricata, Laomedea 1 1 4 
divisa, Campanularia 123 
dubia, Obelia 114 
dubia, Thaumantias 73 
dumosa, Lafoea 47, 65, 1 22 

echinata, Hydractinia 43 
edwardsi, Campanularia 73, 81 
edwardsi, Clytia 8 1 
edwardsia, Clytia 43 
elongata, Campanularia 107, 110 
elsaeoswaldae, Clytia 80 
Epenthesis 70, 71 

cymbaloidea 73, 74 

mccradyi 87 

equilateralis, Obelia 1 14 
Eucalix 48-49, 50-51 

morgansi 5 1 

retroflexus 5 1 
Eucampanularia 47-48, 5 1 , 52, 97 

groenlandica 52 

integra 52 



speciosa 52 
verticillata 52 
Eucope 70, 71, 1 12 
affinis 71,73,74 
alternata 1 1 4 
articulata 1 1 4 
campanulata 7 1 , 
diaphana 1 14 
exigua 73 
fusiform is 1 14 
gemmifera 73 
pamsitica \ 1 4 
pictall, 123 
polygena 1 14 
polystyla 1 1 
primordiale 123 
pyriformis 1 1 4 
thaumantias 74 
thaumahtoides 71 
variabilis 73 

73, 74 


bilabiata 58 

caliculata 58,61 

campanularia 58, 60, 6 1 , 63, 67 


eucophora (& eucopophora), Clytia 123 
Eucopium primordiale 123 
Eulaomeda 97 

flexuosa 105, 117 
Eulaomedea 47, 97-98, 107 

angulata 98 

calceolifera98, 102 

flexuosa 98, 107, 1 17 

pseudodichotoma 1 1 1 
everta, Campanularia 51,58, 60 
<?v<?/7a, Obelia 1 14 
, Orthonia 5 1 
, Orthopyxis 60 
exigua, Eucope 7 3 
exigua, Laomedea 102, 103, 104, 105 

fabredomergui, Obeliopsis 120 

fastigiata(um), Campanularia & Stegopoma 123 

flabellata, Campanularia 1 14 

flabellata, Obelia 1 14, 120 

flavidula, Clytia 73, 74, 76, 77, 80, 81 

.flavidula, Oceania 73, 81 

flemingii, Campanularia 95, 96, 1 1 3 

flexuosa, Campanularia 105 

flexuosa, Eulaomeda 105, 1 1 7 

flexuosa, Eulaomedea 98, 107, 1 1 7 

flexuosa, Laomedea 40, 41 42 43 47 94 

105-107; Fig. 19, p. 105 
folleata, Clytia 91 
folleatum, Phialidium 9 1 
forbesi, Thaumantias 74 
./&*/. Clytia 84, 86 
fragilis, Campanularia 107, 109, 110 

fragilis, Obelia 1 14 
fruticosa, Campanularia 121, 122 
fruticosa, Lafoea 122 
fruticosa, Laomedea 1 24 
fruticosa, Sertularia 121, 124 
fruticosus, Thyroscyphus 121 
fusiformis, Eucope 1 14 

gardineri, Clytia 91 

gardineri, Phialidium 9 1 

Gastroblasta 48-49, 70, 72 
raffaelei 74, 81 

gaussi, f. of Obelia geniculata 1 14 

gegenbauri, Campanularia 73, 80 

gelatinosa, Campanularia 95 

gelatinosa, Hartlaubella 40, 43, 44 48 95-97 
113, 115, 118,119; Fig.16, p. 92 

gelatinosa, Laomedea 47, 95, 107 

gelatinosa, Obelaria 95 

gelatinosa, Obelia 95 

gelatinosa, Sertularia 48, 94, 95 

gemmifera, Eucope 73 

genicolata, Sertolare 1 1 4 

geniculata (sens. Lister), Campanularia 9 1 

geniculata, Clytia 84, 86 

geniculata, Monopyxis 1 1 3 

geniculata, Obelia 40, 42, 43, 45, 96 101-102 

geniculata, Sertularia 1 12, 1 13, 1 14, 1 19 

gigantea, Campanularia 73, 75, 81 

gigantea, Clytia 40, 74, 77 

gigantea, Laomedea 74 

Gonothyrea 9 1 

Gonothyraea 47, 48, 49, 91-92 
/mz//mz92,93,94, 119 
longicyatha 1 1 4 
loveni 40, 41, 42, 43, 44 49 73 79 91 

92-94; Fig. 15, p. 92 
gracilis, Campanularia sens. Allman 60, 61 , 66; 

sens. Meyen 50; sens. Sars 74, 77, 8 1 
gracilis, Clytia 42, 7 1,91 
gracilis, Laomedea 4 1 , 7 1 , 73, 74, 75, 76 78-79 

91,92,94, 114 
gracilis, Lomedea 78, 117 
gracilis, Obelia sens. Calkins 1 14 
gracilis, Silicularia 50, 118-119 
gracillima, Campanularia & Lafoea 122 
Grammaria abietina 122 
grandis, var. of Rhizocaulus verticillatus 69 
grandis, Stegella 69 
gravieri, Campanularia 84, 85 
gravieri, Clytia 42, 85, 86 
griffini, Obelia 1 14 

groenlandica, Campanularia 52, 55, 56, 57 
groenlandica, Eucampanularia 52 
gymnopthalma, Obelia 1 14 

Hartlaubella 48-49, 9 1,94 
gelatinosa 40, 43, 44, 48, 95-97, 113, 115, 
118, 11 9; Fig. 16, p. 92 



helgolandica, Obelia 1 14 

hemisphaerica, Clytia 40, 4 1 , 42, 43, 45, 70-7 1 , 

73-82, 86, 87, 89, 90, 9 1 , 92, 94; Fig. 9, 

p. 75 

hemisphaerica, Medusa 70, 71, 73, 74 
hemisphaerica, Oceania 73, 78 
hemisphaerica, Thaumantias 73, 74 
hemisphaericum, Phialidium 7 1 , 74, 77 
hendersonae, Clytia 84 
hexaradiata, f. of Pseudoclytia pentata 74 
hicksoni, Campanularia 54 
Hincksella 123 
Hincksia 57, 58 

tincta 58 
hincksii, Campanularia 5 1 , 53-55, 80, 82; 

Fig. 3, p. 55 
hincksii, Paracalix 5 1 
humilis, Campanularia & Cuspidella 123 
hummelincki, Campanularia 82 
hummelincki, Clytia 41, 42, 44, 69, 82-83, 86; 

Fig. 10, p. 82 

hummelincki, Laomedea 82 
hyaliana, Obelia 92, 1 14, 1 17, 119 
hyalina, Gonothyraea 92, 93, 94, 1 19 
hyalina, Obelia 92, 94, 1 14, 119 
hybridum, var. of Staurostoma laciniatum 84 
Hydractinia echinata 43 
atlantica 50 
repens 50, 66 
Hypanthia 50 

inconspicua, Campanularia 73 

inconspicua, Thaumantias 73, 74, 76 

indivisa, Tubularia 123 

integra, Campanularia 52, 60 

integra, Capsularia 6 1 

integra, Eucampanularia 52 

integra, Orthopyxis 40, 42, 44, 48, 57, 60-67, 

78, 122, 123; Fig. 6, p. 62 
integriformis, Campanularia 6 1 , 64, 66-67 
intermedia, Campanularia 58, 60 
intermedia, f. of Obelia geniculata 1 14 
intermedia, f. of Orthopyxis ( = Campanularia) 

crenata 59 

intertexta, Campanularia 65, 122 
intertexta, Conchella 122 
irregular is, Obelia 1 14 
islandica, Clytia* 42 , 84; Fig. 1 1 , p. 84 
islandicum, Phialidium 84 

johnstoni, Campanularia 7 1 , 73, 75, 77, 79-80 

johnstoni, Clvthia 74 

johnstoni, Clytia 69, 70, 73, 74, 80 

kincaidi, Campanularia, Clytia, Laomedea & 

Obelia 91 
Kirchenpaueria pinnata 124 

lacerata, Calycella, Campanularia, Capsularia 

& Laomedea, 122 
lacerata, Opercularella 1 10, 122 
lacerata, Wrightia 122 
laciniatum, Staurostoma 84 
laevis, Campanularia 54, 60, 6 1 , 65 
laevis, Capsularia 6 1 , 65 
/aev/s,dytia55,57, 73 
Lafoea 122 

dumosa47,65, 122 

fruticosa 122 

gracillima 122 

parvula 122 
lairii, Laomedea 1 44 
languida, Clytia 8 1 
languida, Oceania 74 
languidum, Phialidium 74, 81 
Laomedea 41,47,48,49,91, 97-98, 1 22 

acuminata 122 

angulata 41, 42, 43, 44, 98-102, 104, 105, 
125; Fig. 17, p. 99 

bicuspidata var. picteti 1 1 4 

bicuspidata var. tennis 1 14 

brochi 47, 107,110 

calceolifera 41, 42, 43, 44, 47, 98, 100, 101, 

102-105; Fig. 18, p. 103 
conferta4\, 102-103, 104 
congdoni 1 14, 119 
decipiens 107 ', 109,110 
divaricata 1 1 4 
exigua\02, 103, 104, 105 
flexuosa 40, 41, 42, 43, 47, 94, 97, 105-107; 

Fig. 19, p. 105 
fruticosa 124 
gelatinosa47,95, 107 
gigantea 74 
gracilis 41, 71, 73, 74, 75, 76, 78-79, 91, 92, 

94, 114 

hummelincki 82 
kincaidi 9 1 
lairii 1 14 

lautalQl, 108,110 
neglecta 40, 43, 47, 48, 96, 107-111; 

Fig. 20, p. 109 
obliqua 122 

plicata \\1.\\9 

pseudodichotoma41,44, 111-112; Fig. 21, 
p. Ill 

repens 60, 6 1 , 62, 66 
sargassi 1 14, 119 
sauvagii 121 
sphaeroidea 98, 102 
spinulosa 1 14 
tenuis 122-123 
lauta, Laomedea 107, 108, 110 



laxa, Sertularia 121 
lennoxensis, Campanularia 58, 60 
Leptomedusa 57,61 
Leptoscyphus tennis 122-123 
leucostyla, Obelia 96 
leucostvla, Thaumantias 74, 96 
linearis,* Clytia 84-86; Fig. 12, p. 85 
linearis, Obelia 84 
lineata, Thaumantias 74 
Lomedea 97 

gracilis 78, 1 1 7 
longa, Obelia 1 1 4 
longicyatha, Clytia 120 
longicyatha, Gonothyraea 1 1 4 
longicyatha, Obelia 1 14, 120 
longissima, Obelia 40, 4 1 , 96, 1 1 4, 1 1 8 
longissima, Sertularia 1 1 4 
longitheca, Clytia & Obelia 1 14 
loveni, Gonothyraea 40, 41, 42, 43, 44, 49, 73, 

79, 9 1,92-94; Fig. 15, p. 92 
loveni, Laomedea 9 1 , 92, 97 
loveni, Obelia 92 
lucida. Medusa 72, 73 
lucida, Thaumantias 73 

madeodi, Plumularia 124 

macrogona, Orthopyxis 6 1 

macrotheca, Campanularia 53, 54 

magnifica, Campanularia 52, 53 

maior, Campanularia 1 14 

marina, Medusa 1 1 2 

mccradyi, Clytia* 42, 87-88; Fig. 13, p. 87 

mccradyi, Epenthesis, Oceania & Phialidium 87 

Medusa 4%, 10,11,124 

campanella 73 

cymbaloidea 73 

hemisphaerica 70, 71, 73, 74 

lucida 72, 73 

marina 1 12 

microtheca, Obelia 1 14 
minor, var. of Laomedea spinulosa 1 14 
mira, Agastra 6 1 , 67 
Modeeria rotunda 123 
Monopyxis 1 12, 113 

geniculata 1 1 3 
Monosklera4&, 112 

pus ilia 1 14 

Monotheca obliqua 113, 122 
morgansi, Campanularia & Eucalix 5 1 
multidentata, Obelia 1 14 
Multioralis 70, 72 
mutabilis, Campanularia 123 
mutabilis, Scandia 123 

neglecta, Campanularia 107 
neglecta, Laomedea 40, 43, 47, 48, 96, 107-111; 
Fig. 20, p. 109 

neglecta, Paralaomedea 47, 98, 107 
Nemertesia antennina 1 2 1 

comata 121 

ramosa 121, 122 
nigrocaulus, Obelia 1 14 
nodosa, Obelia 1 14 
noliformis, Campanularia 80 
noliformis, Clytia 73, 80, 81 
novaezealandiae, Billardia 123 


Obelia 39, 41, 42, 47, 48, 49, 77, 91, 94, 96, 97, 

112-113, 124 
adelungi 1 14 
adriatica 120 
alternata 1 1 4 
andersoni 1 14 
angulosa 1 14 
arruensis 1 1 4 
attenuata 1 14 
aus tralis 1 14 
austrogeorgiae 113, 117 
bicuspidata9Q,113, 114 
bidentata 40, 43, 44, 85, 96, 97, 110, 111, 

bifurcata 1 14 
biserialis 114 
borealis 1 1 4 
braziliensis 1 1 4 
calceolifera 102 
chinensis 1 14 

commensuralis 114, 1 17, 118 
commissuralis 114, 1 17, 119 
congdoni 1 14, 119 
corona 114 
coughtreyi 114 
dichotoma 40, 41, 43, 45, 73, 96, 100, 101, 

102, 104, 112, 113, 114, 117-119, 120, 


dischotoma 1 14, 117 
dubia 1 1 4 
equilateralis 1 1 4 
everta 1 1 4 
flabellata\\4, 120 
fragilis 1 1 4 
gelatinosa 95 
geniculata 40, 42, 43, 45, 96, 101-102, 113, 

gracilis 1 14 
griffini 1 14 
gymnopthalma 1 14 
helgolandica 1 1 4 
hyaliana92, 114, 117, 119 
hyalina 92, 94, 114,119 
irregularis 1 1 4 
kincaidi 9 1 



leucostyla 96 

linearis 84 

longa \ \ 4 

longicyatha 1 14, 120 

longissima 40, 4 1 , 96, 1 1 4, 1 1 8 

longitheca 1 1 4 

loveni 92 

microtheca 1 1 4 

multidentata \ 14 

nigrocaulus 1 14 

nodosa 1 1 4 

obtusidentata 1 1 4 

oxydentata \ 1 4 

paulensis 88, 90 

piriformis 1 1 4 

plana 120 

p//cflta40,43, 114, 117, 119 

polystyla 1 1 

pygmaea 1 1 4 

pyriformis 1 14 

racemosa \ \ 4 

rhunicola 1 14 

solowetzkiana 1 14 

sphaerulina 1 12 

spinulosa 113, 114 

surcularis 1 1 4 

tennis 1 14 

undotheca 1 1 4 
obeliformis, Clytia 73, 75, 8 1 
Obeliopsis\l2, 113 

fabredomergui 1 20 
ObelissaW, 112 
Obelomma 112,113 
obliqua, Campanularia 84 
obliqua, Clytia 85, 86 
obliqua, Laomedea 1 22 
obliqua, Monotheca 113, 122 
obliqua, Plumularia 113, 122 
obtusidens, Campanularia 1 14 
obtusidentata, Obelia 1 14 
Oceania 70, 71,87 

armata 8 1 

discoida 72 

hemisphaerica 73, 78 

languida 74 

mccradyi 87 
olivacea, Clytia 67 
Opercularella lacerata 110, 122 

everta 5 1 

Orthopyxis 4 1 , 48, 49, 50, 5 1 , 57-58 

africana 39 , 60 

asymmetrica 6 1 , 67 

caliculata 48, 6 1 , 63, 65-66 

clytioides 50, 118-119 

compressa 6 1 , 66 

crenata 40, 42, 52, 58-60; Fig. 5, p. 59 

delicata 58-59, 60 

everta 60 

//iteffra 40, 42, 44, 48, 57, 60-67, 78, 122, 

123; Fig. 6, p. 62 
macrogona 6 1 
poterium 57-58, 60 
tincta* 58 

volubiliformis 57, 74 
oxydentata, Obelia 1 14 


hincksii 51 

pulcratheca 5 1 

volubilis 51 
Paralaomedea 47, 97, 98 

brochii 47 

neglecta47,98, 107 
parasitica, Eucope \ 14 
parvula, Campanularia 122 
parvula, Lafoea 122 
paulensis, Campanularia 88 
paulensis, Clytia 42, 43, 44, 76, 85, 86, 88-91, 

113, 11 5, 117; Fig. 14, p. 89 
paulensis, Obelia 88, 90 
pelagica, Campanularia 73, 78-79 
pelagica, Clytia 73 
pelagica, Laomedea 74, 75-76, 8 1 
pentata, Clytia 42, 91 
pentata, Phialidium 9 1 
pentata, Pseudoclytia 74 
Phialella quadrata 123 
Phialidium 48, 69, 70, 71, 72 

bicophorum 74, 80 

buskianum 74 

discoidum 73 

folleatum 9 1 

gardineri 9 1 

hemisphaericum 1 1 , 74, 77 

islandicum 84 

languidum 74, 81 

mccradyi 87 

pentata 9 1 

temporarium 74 

variabile69, 74 

Phialium 69 
picta, Eucope \ 1 4 

picteti, var. of 'Laomedea bicuspidata '114 
pileata, Thaumantias 74 
pinnata, Kirchenpaueria 124 
piriformis, Obelia 1 14 
plana, Obelia & Thaumantias 120 
platycarpa, Campanularia 61 



Platypyxis 70,71,80 

cylindrica 73 

plicata, Laomedea 117, 119 
plicata, Obelia 40, 43, 1 14, 1 1 7, 119 
Plumularia macleodi 124 

obliqual!3, 122 
Podocoryne carnea 43 
polygena, Eucope 1 1 4 
polystyla, Eucope 1 1 
polys tyla, Obelia 71 
posterior, Clytia 6 1 
poterium, Clythia, Clytia & Orthopyxis 57-58, 

primordiale, Eucope 123 
prolifera, Campanularia 1 14 
Pseudoclytia 70, 72 

pentata f. hexaradiata 74 
pseudodichotoma, Eulaomedea 1 1 1 
pseudodichotoma, Laomedea 41, 44, 111-112; 

Fig. 21, p. Ill 

ptychocyathus, Campanularia 80 
pulcratheca, Campanularia 5 1 
pulcratheca, Paracalix 5 1 
punctata, Thaumantias 74 
pusilla, Monosklera 1 14 
pygmaea, Obelia 1 14 
pyriformis, Eucope 1 14 
pyriformis, Obelia 1 14 

quadrata, Phialella 123 

racemosa, Obelia 1 14 

raffaelei, Gastroblasta 74, 81 

ramosa, Cymodocea 121 

ramosa, Nemertesia 121, 122 

ramosum, Schizocladium 1 1 4 

rara, Campanularia 53 

raridentata, Campanularia 73, 74, 75, 76, 80, 


raridentata, Thaumantias 74 
repens, Hypanthea 50 
repens, Laomedea 60, 6 1 , 62, 66 
repens, Silicularia 50 
retroflexa, Campanularia 5 1 
retroflexus, Eucalix 5 1 
Rhizocaulus 41 , 48, 49, 50, 52, 67 

verticillatus 40, 4 1 , 43, 48, 52, 57, 67-69; 
Fig. 7, p. 68 

verticillatus var. grandis 69 
rhunicola, Obelia 1 14 
rijckholtii, Clytia 6 1 , 65 
ritteri, Campanularia 60 
rosea, Silicularia 50 
rotunda, Modeeria 123 
rubra, Agastra6l,63,61 
ryckholtii, Clytia 6 1 , 65, 1 23-124 

sargassi, Laomedea 114, 119 
sarnica, Thaumantias 74 

sarsi, Clytia 4 1,74, 78 
sauvagii, Laomedea 121 
Scandia mutabilis 123 
Schizocladium 48, 112 

ramosum 1 14 

sericea, Sertularia [Bryozoa] 96 
serrata, Clytia 85 
serrulata, Campanularia 73, 80 
serrulata, Clytia 73, 74, 76 
Sertolare genicolata 1 14 
Sertularia 70-7 1 

dichotoma 1 1 7 

fruticosa!21, 124 

gelatinosa48,94, 95 

geniculata 1 12, 1 13, 1 14, 1 19 

laxa 121 

longissima 1 14 

(Monopyxis) geniculata 1 1 3 

sericea [Bryozoa] 96 

spinosa [Bryozoa] 96 

syringa 70, 78 

uniflora 55, 57, 73, 74, 77-78 


volubilis 48, 5 1 , 52, 55, 70, 73, 77-78 
Silicularia 48, 49, 50, 57,70 

atlantica 50 

bilabiata 60 

clytioides 50, 118-119 

gracilis 50, 118-119 

repens 50 

rosea 50 

simplex, Clytia 73, 8 1 
simplex, Cymodocea 114, 1 17, 121 
Slabberia 112 
solowetzkiana, Obelia 1 14 
speciosa, Campanularia 52, 53 
speciosa, Eucampanularia 52 
speciosus, Campanularia 52 
sphaeroidea, Laomedea 98, 102 
sphaerulina, Obelia 1 12 
spinosa, Sertularia & Vesicularia [Bryozoa] 96 
spinulosa, Campanularia 1 14 
spinulosa, Laomedea 1 14 
spinulosa, Obelia 1 13, 1 14 
Staurophora 84 
Staurostoma 84 

laciniatum 84 
Stegella grandis 69 
Stegopomafastigiatum 123 
striata, Clytia 86 
striata, Obelia 85 

subantarctica, f. of Obelia geniculata 1 14 
subsessilis, f. of Obelia geniculata 1 14 
subtropica, f. of Obelia geniculata 1 14 
surcularis, Obelia 1 14 
syringa, Calycella 55, 57, 78, 122 
syringa, Sertularia 70, 78 

temporarium, Phialidium 74 



tenuis, Campanulina, Laomedea & Lepto- 

scyphus 122-123 

tenuis, var. of 'Laomedea bicuspidala' 1 14 
tenuis, Leptoscyphus 122-123 
tenuis, Obelia 1 14 
thaumantias, Eucope 74 
Thaumantias 4%, 1Q,1\, 112 

buskiana 74 

convexa 73 

diaphana 1 14 

dubia 73 

forbesi 74 

hemisphaerica 73, 74 

inconspicua 73, 74, 76 

leucostyla 74, 96 

lineata 74 

lucida 73 

pileata 74 

plana 120 

punctata 74 

raridentata 74 

sarnica 74 


typica 74 

thaumantoides, Eucope 71,73 
thompsoni, Thaumantias 73, 74 
Thyroscyphus fruticosus 1 2 1 
tincta, Campanularia 58, 60 
tincta, Hincksia & Orthopyxis* 58 
Trochopyxis 70 

bicophora 73, 80 (as Clytia) 
Tubularia clytioides 50, 1 14, 1 17, 118 

indivisa 123 

tulipifera, Campanularia & Tulpa 70 

tulipifera & tulpifera 70 
tulpifera, Tulpa 70 
typica, f. of Laomedea angulata 100 
typica, Thaumantias 74 

. Clytia 88, 91 
undotheca, Obelia 1 14 
undulata, Clytia 60-6 1 , 65 
uniflora, Clytia 74 

uniflora, Sertularia 55, 57, 73-74, 77-78 
urnigera, Clytia 64-65, 73, 78 

var iabile, Phialidium 69, 74 

\ariabilis, Eucope 73 

vermicularis, Campanularia 120 

verticellata, Campanularia 67 

verticillata, Campanularia 67, 69 

verticillata, Campanulata 68 

verticillata, Eucampanularia 52 

verticillata, Sertularia 48, 5 1-52, 67, 70 

verticillata, Verticillina 68 

verticillatus, Rhizocaulus 40, 41 , 43, 48, 52, 57, 

67-69; Fig. 7, p. 68 
Verticillina 67 

verticillata 68 

Vesicularia spinosa [Bryozoa] 96 
vicophora, Clytea & Clytia 73 
villafrancensis, Campanularia 74, 8 1 
villafrancensis, Clytia 74 
viridicans, Clytia 74 
viridicans, Phialidium 71 , 73, 74, 86 
vitrina, Aequorea 123 
volubiliformis, Campanularia 73, 80 
volubiliformis, Orthopyxis 57, 74 
volubilis, Campanularia 48, 51, 54, 55-57, 69, 

70, 73, 76, 77, 79; Fig. 4, p. 55 
volubilis, Clytia 73, 74, 76, 77, 78 
volubilis, Eucampanularia 52 
volubilis, Paracalix 5 1 
volubilis, Sertularia 48, 51, 52, 55, 70, 73, 

volubis, Clytia 43 

Wrightia 122 

British Museum (Natural History) 
Chance , change & challenge 

Two multi-author volumes from one of the foremost scientific institutions in the world. 
General Editor: P. H. Greenwood 

The Evolving Earth 

Editor: L. R. M. Cocks 

The Evolving Biosphere 

Editor: P. L. Forey 

In the first volume, The Evolving Earth, twenty scientists have been asked to review 
the present state of knowledge in their particular field, ranging from the origin of 
the Earth, through ocean sediments and soils to continental drift and palaeogeography. 

In the companion volume, The Evolving Biosphere, museum scientists have chosen 
an evolutionary concept speciation, coevolution, biogeography etc. and related 
this to the group of animals or plants in which they are specialising. Thus beetles 
and birds exemplify sympatric and allopatric speciation, butterflies mimicry and 
certain fishes explosive evolution. 

In both volumes the text is supplemented by over one hundred specially-commissioned 
pieces of two-colour artwork. 

These two books will be invaluable to all sixth-form and undergraduate biology and 
geology students. 

The Evolving Earth: 276x219 mm, 280pp, 138 line illustrations, 42 halftones 
The Evolving Biosphere: 276x219 mm, approx. 320pp, 133 line illustrations 
Published: May 1981 

Co-published by the British Museum (Natural History), London and Cambridge 
University Press, Cambridge. 

Titles to be published in Volume 42 

The tick collection (Acarina: Ixodoidea) of 

the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and 

general collections of the British Museum (Natural History). 

By James E. Keirans 

Hydroids and medusae of the family Campanulariidae recorded 
from the eastern North Atlantic, with a world synopsis of genera. 
By P. F. S. Cornelius 


Printed by Henry Ling Ltd, Dorchester 


Bulletin of the 

British Museum (Natural History) 


Zoology series Vol 42 No 3 27 May 1982 

The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and 
an Historical series. 

Papers in the Bulletin are primarily the results of research carried out on the unique and 
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specialists from elsewhere who make use of the Museum's resources. Many of the papers are 
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Parts are published at irregular intervals as they become ready, each is complete in itself, 
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 

Trustees of the British Museum (Natural History), 1982 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology : Dr J. G. Sheals 
Editor of Bulletin : Dr C. R. Curds 
Assistant Editor : Mr C. G. Ogden 

ISSN 0007-1498 Zoology series 

Vol 42 No 3 pp 149-225 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 27 May 1982 




A new species of free-living nematode from the Firth of Clyde, Scotland. By P. J. D. 

Lambshead 149 

Notes on Atlantic Asteroidea 2. Luidiidae. By A. M.Clark 157 

New and little known species of Oncaeidae (Cyclopoida) from the Northeastern 

Atlantic. By S. J. Malt 185 

Larval and post-larval development of the Slender-legged Spider Crab, Macropodia 
rostrata (Linnaeus), (Oxyrhyncha: Majidae: Inachinae), reared in the laboratory. 
By R.W. Ingle 207 

A new species of free-living nematode from the 
Firth of Clyde, Scotland 

P. J. D. Lambshead 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 


A new species of free-living marine nematode, Odontophoroides paramonhystera is 
described from a fine sandy beach in the Firth of Clyde, Scotland. It is similar to 
Odontophoroides monhystera (Gerlach, 1953) from which it differs primarily in having a 
precloacal supplement. Odontophoroides monhystera sensu Hopper, 1968, is renamed 
Odontophoroides hopperi nom. nov. 

Odontophoroides paramonhystera sp. nov. 
Figs 1,2, 3 &4 

MATERIAL STUDIED. Eleven males and nine females, deposited at the British Museum 
(Natural History): Holotype- d 1 ,, BM(NH)198 1.6.42; Paratypes- d 1 , BM(NH) 
1981.6.43-50 & 1981.6.59-60,$,^ BM(NH)198 1.6.5 1-57 & 1981.6.61-62. Also an 'en face' 
view of a juvenile was prepared, BM(NH)1981.6.58. In addition specimens of O. 
monhystera, collected by Dr S. Lorenzen (1973) and Drs G. Boucher & M. N. Helleouet 
(1977), were examined. 

LOCALITY. Low water spring mark, fine sandy beach, Scalpsie Bay, Isle of Bute, Scotland. 
MEASUREMENTS. See Table 1 . 

DESCRIPTION. Slender, colourless, worms. No pigment spots or ocelli seen in preserved 
specimens. Cuticle finely, transversely striated (Fig. 3a, b). Six papillae visible in 'en face' 
view (Fig. Ic, juvenile): whether R, or R 2 not clear. Four R 3 cephalic setae, about 1J head 
diameters long. Two fine subcephalic setae associated with the subventral R 3 cephalic setae 
in females; not observed in males. Six cervical setae present; two subdorsal and four sub- 
ventral, posterior subventral cervical setae more posterior in males than females. Stout 
lateral setae at base of amphids; about 15 //m long, but difficult to measure accurately due to 
the angle at which they project. Somatic setae in four irregular rows; length variable, but 
often longer in oesophagus and anterior gut regions. Amphids prominent, shaped like the 
head of a looped Shepherd's crook, with cuticularized borders; wider in males. Corpus 
gelatum displaced in about half of the male specimens but none of the females. Buccal cavity 
large, conical, cuticularized, with a complex arrangement of teeth (Fig. Ic, d). Six teeth 
present, bifurcate with a minor projection growing out of the main tooth. The four sublateral 
teeth are large and heavily cuticularized. Lateral teeth reduced to flat plates. Teeth attached 
at base to heavily cuticularized anterior section of buccal cavity (Fig. Id). Two subdorsal, 
cuticularized tooth-like projections present; no equivalent subventral structures. In some 
preserved specimens the mouth was open, partially everting the teeth. Oesophagus 
muscular, especially in region which surrounds posterior section of buccal cavity; bulbs 
absent. Excretory pore prominent, located opposite posterior half of buccal cavity. Cellular 
sac in close association with posterior of excretory cell (Fig. 2b); function unknown. Caudal 
glands present lying entirely posterior to the anus. Tail conical. 

Bull. Br. Mus. nat Hist. (Tool.) 42 (3): 149-155 Issued 27 May 1982 







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Fig. 1 (a) Head of male; (b) head of female; (c) 'en face' view of juvenile (dorsal is to the right of 
illustration); (d) teeth of female (dorsal is to the right of illustration); (e) cloacal region. Bar 

scale = 1 5 yum. 

Male. Spicules paired, equal, arcuate, cephalate at proximal end slightly hooked at distal 
end; about 4 a.b.d. long. Gubernaculum appears to surround spicules and has narrow, 
paired, dorso-caudally directed apophyses. One supplement, with pore, 1 a.b.d. anterior to 
cloaca. Two testes, not clearly visible, outstretched, in tandem, ventral and apparently to left 
of gut; posterior testis about 250 //m behind anterior testis. Vas deferens widens into sac-like 
tube, of granular appearance, 4 a.b.d. anterior to cloaca. 

Female. Vulva in anterior position. Vagina with cuticularized walls, surrounded by band 
of muscle, which is attached to ventral cuticle, the uterus, and to a tube-like structure. This 
tubed structure can give the false impression of connecting the sac associated with the 
excretory cell to the vagina (Fig. 2b). Muscle band presumably controls the opening of the 



Fig. 2 (a) Whole female; (b) vulval region; (c) tail of male. Bar scale = 1 5 //m. 

vulva. Single posterior ovary, outstretched, extending for most of the length of the body, 
ventral and to left of gut. 

DIFFERENTIAL DIAGNOSIS. Odontophoroides paramonhystera sp. nov. is very similar to O. 
monhystera (Gerlach, 1953): the females and juveniles appear morphologically 
indistinguishable. However, males of the new species possess a precloacal supplement 
(lacking in O. monhystera) and the spicules are distally less slender. 



Fig. 3 Anterior of female, left lateral views: (a) showing amphid, cephalic and subcephalic setae, 
and lateral seta; (b) showing fine subcephalic seta associated with subventral R 3 seta. 


The previously monotypic genus Odontophoroides was erected by Boucher & Helleouet 
(1977) for Synodontium monhystera Gerlach, 1953. Gerlach (1953) originally described S. 
monhystera from Italy, and the types have been lost (Riemann, pers. comm.). However 
Gerlach (1953) noted 'Praanal-organe fehlen'. Subsequently a female was described by 
Hopper (1968) from Prince Edward Island, Canada, and males, females and juveniles 
described by Lorenzen (1973) from Helgoland, North Sea. Boucher & Helleouet (1977) also 
described a male (which has unfortunately been lost, Boucher pers. comm.), females and a 
juvenile from the English Channel. The type material of S. monhystera by implication, 
remains the specimens described by Gerlach (1953). Gerlach's (1953) and Boucher & 
Helleouet's (1977) descriptions show the distal ends of the spicules as less slender than 
Lorenzen's (1973) specimens. This character may, therefore, prove to be unreliable. 

Odontophoroides monhystera sensu Hopper, 1968, is depicted without lateral subcephalic 
setae. The text refers to two lateral subcephalic setae, but notes them as positioned '8 // and 
23 //, respectively, from anterior extremity' which would not seem to place them in their 
typical situation, i.e. at the base of the amphids. Lorenzen (1973) demonstrates that such 
setae are only present in adults and fourth stage larvae and Boucher (1974) shows the range 
of variation which may occur in the position of subcephalic setae in the related genus 
Odontophora, where species may or may not have lateral subcephalic setae. The presence or 
absence of these setae may therefore not be a stable generic character. In other respects 
Hopper's (1968) specimen is clearly Odontophoroides, being monodelphic (V= 19%) and 



Fig. 4 (a) Left lateral view of female head showing subventral and subdorsal teeth, and subdorsal 
tooth like projection; (b) right lateral view of female head showing lateral tooth. 

having 'six prominent bifid odontia'. As females of this genus appear indistinguishable it 
cannot safely be referred to either species and so to avoid zoogeographic problems it is here 
renamed Odontophoroides hopperi nom. nov. (type Synodontium monhystera sensu 
Hopper, 1968). It is characterized by the lack of lateral subcephalic setae at the base of the 
amphid and the absence of fine subcephalic setae associated with the subventral R 3 
subcephalic setae. The discovery of further specimens, particularly males, should elucidate 
the relationship between O. hopperi and the European species but until such time the status 
of this species remains equivocal. As a consequence of this analysis the original generic 
diagnosis of Odontophoroides given by Boucher & Helleouet (1977) should be modified to 
include forms with or without lateral subcephalic setae. 


I would like to express my thanks to Drs Riemann, Lorenzen and Boucher for kindly lending 
specimens of O. monhystera, to Dr Jensen for helpful discussion, to Mr Coles for technical 
advice and to the BM(NH) Photographic Unit for the photomicrographs. 


Boucher, G. 1974. Nematodes libres marins des lies Hautes de Polynesie. 1-Comesomatidae et 
Axonolaimidae. Cah. Pacif. 17 : 205-232. 


Boucher, G. & Helleouet, M. N. 1977. Nematodes des sables fins infralittoroux de la Pierre Noire 

(Manche occidentale). III. Araeolaimida et Monhysterida. Bull. Mus. natn. Hist. nat. Paris 427, 

Zoo/. 297: 85-122. 
Gerlach, S. A. 1953. Die Nematodenbesiedlung des Sandstrandes und des Kustengrandwassers an der 

Italienischen Kiiste. I. Systematischer Teil. Arch. Zoo/. Ital. 37 : 517-640. 
Hopper, B. E. 1968. Marine nematodes of Canada. I. Prince Edward Island. Can. J. Zoo/. 

46: 1103-1111. 
Lorenzen, S. 1973. Freilebende Meeresnematoden aus dem Sublitoral der Nordsee und der Kieler 

Bucht. Veroff.Inst. Meeresforsch. Bremerh. 13 : 307-316. 

Manuscript accepted for publication 8 June 1981 

Notes on Atlantic Asteroidea. 2. Luidiidae 

Ailsa M. Clark 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 


In reviewing the Atlantic species of Asteroidea, recent proposals for inclusion of the family 
Luidiidae in the order Paxillosida rather than the Platyasterida (as in the 'Treatise of 
Invertebrate Paleontology') need to be discussed. Also, several taxonomic complications to 
do with the large genus Luidia have become apparent, particularly in reassessing the ranks of 
similar taxa from the continental shelves on the two sides of the ocean. These are better 
disposed of in a preliminary paper. 

The large collections of the Pillsbury and Gerda of the University of Miami from both 
sides of the tropical Atlantic have supplemented the material already in the British Museum 
(Natural History). Only L. patriae Bernasconi from Argentina is not available, so comments 
on its affinity with L. scotti Bell are based on the published description. The number of 
nominal species occurring in the Atlantic is reduced by synonymy or lowering to infra- 
specific rank from 17 to 11. Coverage ofL. atlantidea Madsen, L. barbadensis Perrier, L. 
ciliaris (Philippi) and L. senegalensis (Lamarck), all of which are limited to either east or 
west, is restricted to inclusion in a tabular key of principal characters and an outline of the 
distribution. Also L. dathrata (Say) raises only a nomenclatural problem, being another 
species of limited range. A neotype for L. alternata alternata (Say) and a lectotype for L. 
sarsi sarsi (Diiben & Koren) are designated and discussed. 

The good size range of specimens from the Miami collections has allowed for some 
comments on post-metamorphosal growth changes. 

Systematic account 
LUIDIIDAE Sladen, 1889 

Astropectinidae: Luidiinae Sladen, 1889 : 244. 

Luidiidae: Verrill, 1899 : 201; Fisher, 1911 : 105; Bernasconi, 1943 : 2-3; Fell, 1963 : 433; Spencer & 

Wright, 1966 : U42; Downey, 1973 : 21; Blake, 1973 : 22-24 (pt); A. M. Clark & Courtman-Stock, 

1976 : 43; McKnight, 1977 : 1 16; Blake, 1982 : 186. 

AUTHORITY. Under Article 36 of the International Code of Nomenclature (coordinate 
categories), the authority and date of a name in the family group can originate from its use as 
a subfamily name, so the citation of Verrill, 1899 as author and date for Luidiidae (e.g. by 
Spencer & Wright in the 'Treatise on Invertebrate Paleontology', 1966) is incorrect. 

ORDINAL POSITION. In 1963 Fell, followed by Spencer & Wright (1966) split off the 
Luidiidae to the order Platyasterida (otherwise including only the lower Palaeozoic family 
Palasteriscidae) from its longstanding position close to the Astropectinidae because of its 
transversely matching longitudinal series of adambulacral, actinal and inferomarginal plates. 
This has never seemed to me satisfactory in view of the close resemblance of Luidia and 
Astropecten, especially in transverse section of the arms, and I fully agree with Jangoux 
(1975) and McKnight (1977), who dismissed Fell's disposition because of close anatomical 
resemblances between Luidia and certain astropectinids, and the different alignment of the 
ambulacral and adambulacral plates (the Platasteriscidae having only a very shallow 

Bull. Br. Mus. nat. Hist. (Zool.) 42 (3) : 1 57-1 84 Issued 27 May 1 982 

158 A.M.CLARK 

ambulacral groove, unlike any recent asteroids). The Luidiidae should therefore be included 
in the order Paxillosida Perrier, 1884, formerly lumped with Perrier's Valvatida in Sladen's 
larger order Phanerozonia (1889). Blake (1982) has independently reached the same 
conclusion on the basis of his more detailed morphological studies of the various series of 
ossicles, coupled with the absence of any fossil luidiids in pre-Miocene strata. The Luidiidae 
shares with the Astropectinidae not only the paxilliform abactinal plates but also the 
presence of superambulacral plates, the knobbed tube feet with double ampullae and the 
horizontal framework to the body of enlarged marginal plates (though this is not exclusive to 
the Paxillosida). The main differences are the lack of transverse matching of the actinal series 
with the adambulacrals and inferomarginals in astropectinids a character probably not of 
more than familial weight and the reduction and paxilliform nature in luidiids of the 
superomarginals, so that only their position matching the inferomarginals indicates their 
true identity. (In the Clathrata and Alternata groups of species of Luidia, the two lateralmost 
series of abactinal paxillae each side also match the marginals in length and are indis- 
tinguishable from the superomarginal paxillae unless the latter are broadened which 
reaches an extreme development in Luidia (Platasterias) latiradiata (Gray), referred to the 
Clathrata group by Blake (1973); other groups of Luidia species have smaller lateral 
abactinal paxillae.) 

DIAGNOSIS. A family of Paxillosida with 5-1 1 fairly long, gently tapering arms 
(usually described as 'strap-shaped'), not broadening basally into the disc; lower surface 
almost flat (and in preserved specimens often also the upper, owing to collapse of the fine 
paxillar reticulum); median paxillae irregular and smaller than the lateral ones, of which at 
least the two outermost series are arranged in longitudinal series and matched transversely; 
papulae distributed all over the upper side between the abactinal plates, compound at their 
outer ends; superomarginal plates paxilliform, similar in structure to the adjacent abactinal 
plates, though up to twice as long in species where the abactinal paxillae are particularly 
small (only in the subgenus Platasterias of Luidia are the superomarginals considerably 
broader than the lateral paxillae, though still essentially paxilliform); inferomarginal plates 
massive, lateral or lateroventral in position but never conspicuous in dorsal view, each raised 
into a flat-topped keel bearing some large spines, especially near the upper end (the ambitus 
or broadest part of the arm), interspersed with spinelets of varying size, the sides of the keel 
armed with many fine geometrically-arranged fasciolar spinules; a small actinal plate (rarely 
two proximally) intercalated between each inferomarginal and the matching adambulacral 
plate, often acting as the base for a spiniform pedicellaria (in a few species of the Alternata 
group of Luidia there are supernumerary reduced, pedicellaria-bearing actinal plates 
superimposed on the abradial ends of the adambulacral plates); interradial areas compact, 
rarely with more than a few series of actinal plates; adambulacral plates broad and raised into 
a shallow keel, bearing 2-4 large spines along the crest (the fourth, if present, proximal to the 
third), the innermost (furrow spine) compressed, curved and sabrelike; pedicellariae well 
developed, sessile, with 2-, 3- or even 4-valves, absent in a few species or individuals, those 
of the upper side usually nutcracker-like with broad blunt jaws, the marginal and ventral 
ones more elongated and spiniform; internally, superambulacral plates linking the 
ambulacrals and inferomarginals; tube feet pointed or ending in knobs, their internal 
ampullae double. 

REMARKS. Doderlein (1920) monographed Luidia Forbes, 1839, distinguishing the following 

groups and subgenera: 

Clathrata group: subgenera Senegaster and Petalaster Gray, 1 840; 

Alternata group: subgenera Maculaster, Alternaster and Armaster; 

Quinaria group: subgenera Integraster, Denudaster, Penangaster and Quinaster, 

Ciliaris group: subgenus Hemicnemis Miiller & Troschel, 1840 (other names being 


Fell (1963) pointed out that the type species of both Luidia and Hemicnemis is L. ciliaris 

(Philippi) so that Hemicnemis is a synonym. Except for Armaster, which he lumped with 


Alternaster, he perfunctorily raised all the others to generic rank, a move which has failed 
to gain acceptance, notably from Blake (1973) in an exhaustive study of the ossicles of 
luidiids and related asteroids. At the same time, Blake reduced Platasterias Gray, 1.87 1 to the 
rank of a subgenus of Luidia (whereas Fell had included it in the subclass Somasteroidea), 
since P. latiradiata is undeniably more closely related to Luidia clathrata (Say) than 
clathrata is to L. ciliaris. The family is accordingly considered to be monogeneric. 

Where Atlantic taxa of Luidia are concerned, the limits between Doderlein 's main groups 
are particularly blurred. For instance, L. scotti Bell, 1917 (omitted by Doderlein), now 
treated as a subspecies of L. ludwigi Fisher which Doderlein put under the Alternata group, 
loses its bold colour pattern characteristic of that group soon after preservation and must 
have a different pigment to L. alternata. Since L. ludwigi scotti, together with L. armata, also 
lacks the enlarged abactinal spines found in the majority of species of the Alternata group, it 
bridges the gap to the similarly smooth species of the Clathrata group, both groups having 
the two outermost series of abactinal plates matching up with the marginal plates. L. 
barbadensis Perrier has the lateral paxillae only slightly outnumbering the marginals by c. 
12/10, which serves to differentiate it from the two above groups as defined by Doderlein, 
who listed it as 'incertae sedis' under the Alternata group. Its coarse and squarish lateral 
paxillae certainly link it more closely with the Alternata group than with the Quinaria and 
Ciliaris groups with their finer and more numerous paxillae. 

One of the main characters by which at least the Atlantic species of Luidia can be split up 
is the alignment of the inferomarginal plates, whether or not they extend above the ambitus. 
In the Clathrata and Alternata groups they are ventro-lateral (see Fig. la, b) but in the 
Quinaria and Ciliaris groups they extend higher. The difference in alignment is accompanied 
by some difference in armament. With more ventrally-aligned inferomarginals, the largest 
spines are at the ambitus and project horizontally, or nearly so (though if there is more than 
one ambital spine then the uppermost is more or less reduced); the spines below the ambitus 
are appreciably smaller also and are somewhat appressed. When the inferomarginals extend 
on to the upper surface all their spines are erect, the uppermost one (or two) being largest, the 
top one projecting almost vertically upwards in its natural position, at least on those (often 
alternate) plates where it arises from its highest locus. 

Secondly, the number of lateral paxillae corresponding to the superomarginal ones 
whether matching exactly or in excess is another easily determined character, again 
dividing the Clathrata and Alternata groups from the Quinaria and Ciliaris ones, though L. 
barbadensis is an exceptional intermediate, as already mentioned. 

These two characters are therefore the most important in the tabular key (table 1) to the 
Atlantic species, besides the obvious difference in arm number for some species. The 
occurrence of pedicellariae is variable and not reliable for more than local differences at the 
infraspecific level, though their location and the number of valves may be useful. 

ONTOGENY. The reproduction and larval stages of Luidia sarsi were studied by Tattersall & 
Shepherd (1934). An early post-metamorphosal stage of this species is exemplified by the 
diminutive holotype, R 8 mm, of Astrella simplex Perrier, 1882, synonymized with L. sarsi 

Fully metamorphosed specimens of other species taken by the Pillsbury in the tropical 
Atlantic show that all the primary plates, even the small actinal ones, occur from an early 
stage, though the actinals at first lack any armament. 

At R c. 6 mm, the upperside paxillae of each arm consist of a band of somewhat irregular 
median paxillae flanked on either side by two complete regular longitudinal series, the 
outer of which is the superomarginal series and the other the primary lateral series. Almost 
immediately a secondary lateral series begins to develop proximally between the two but 
probably does not become fully developed to the arm tip until R is c. 25 mm, at least in L. 
alternata numidica Koehler. In L. heterozona Fisher, where the lateral paxillae are relatively 
smaller, the secondary lateral series is not initiated until R c. 25 mm and is still not quite 
complete at R 50 mm. At least in species with fairly coarse paxillae, the primary lateral 



Fig. 1 Half arm sections, viewed from the proximal side, of: (a) Luidia clathrata, Pillsbury st. 
652, Re. 120 mm; (b) L. alternata alternata, Pillsbury st. 767, R c. 90 mm; (c) L. sarsi elegans. 
Albatross st. 2 1 77, R c. 115 mm; (d) L. sagamina aciculata, Atlantide st. 6 1 , R c. 90 mm. 

series can usually be recognised by the fact that both its abradial and adradial basal lobes are 
overlain by the opposing lobes of the secondary lateral and outermost median series of 
paxillae respectively (see Fig. la, b, d) but this does not always hold good when the skeleton 
is more delicate. The median paxillae also increase in number to some extent, less so in 
species where they are stouter, as in L. alternata (Say) where the total number of paxillae 
across the arm proximally between the two opposite superomarginal series increases from 9 
or 10 at R 6 mm to only c. 12 or 13 at R 50 mm. There are still only c. 12 in the largest 
specimen of L. alternata alternata studied, R 175 mm, so the tabula of the larger paxillae in 
such species may be markedly broadened. 

With regard to the armament, as usual in newly metamorphosed asteroids, the spinelets 
and future spines are all attenuated, often trifid, with the terminal point prolonged. This is 
still true up to R c. 9 mm, except for the furrow and second adambulacral spines, which are 
simple and unbranched, though still uniform in length with the other spinelets. At R c. 
10 mm, an ambital inferomarginal spine begins to enlarge in L. alternata and the second 
adambulacral spine also increases disproportionately in size, while a few of the more lateral 
paxillae begin to develop a single large central spine. In species with a third adambulacral 



Fig. 2 (a to d) Dorsal views of a proximal part of an arm, showing the upper ends of several 
inferomarginals, the adjacent superomarginals and some abactinal plates, denuded: (a) Luidia 
ludwigi scotti, paratype, Terra Nova st. 42, R 62 mm; (b) L. barbadensis, Gerda st. 239, R c. 
130 mm; (c) L. sarsi elegans (as in Fig. Ic); (d) L. alternata alternata, Pillsbury st. 767, R c. 
15 mm; (e to g) the same views with armament entire (inferomarginal omitted in (f)) of: (e) L. 
ludwigi scotti (as in Fig. 2a); (f) L. heterozona barimae, Pillsbury st. 652, R 200+ mm; (g) L. sarsi 
africana, Namibia, Re. 100 mm; (h) one adambulacral, actinal and inferomarginal, in ventral 
view of L. alternata alternata (as in Fig. Ib); (i) L. heterozona heterozona, Pillsbury st. 24, R 
100+ mm, lateral view of two successive inferomarginals and superomarginals with a few lateral 
paxillae; (j) apex of jaw of L. heterozona barimae (as in Fig. 20; (k) L. sagamina aciculata, (as in 
Fig. Id), two lines of adambulacral, actinal and inferomarginal plates, the one on the left 
denuded. The proximal side is to the right in (a-g), to the left in (h, i & k). 

162 A.M.CLARK 

Table 1 Tabular key to the Atlantic species of Luidia. Entries in brackets signify occasional 

12 34567 8 9 10 11 























ludwigi scotti 





















alternata (2 subspp.) 






















sagamina aciculata 







+ 2/3 




heterozona (2 subspp.) 







+ 3/2,4/3 




sarsi (3 subspp.) 







+/- 4/3,3/2 











+/- 3/2(4) 




clitoris 7(8) 15-18 c (+) 1 + 3-52 -/3 - 

1 . Arm number. 

2. Colour pattern of upper side: 

+ patchy (may fade in long-preserved L. ludwigi scotti). 

uniform or darker along midradial area or lighter along midradial area or lighter along 

superomarginal or adjoining lateral paxillae. 

3. Number of consecutive lateral paxillae corresponding to 10 superomarginal ones. 

4. Paxillar armament: 

c central spinelets distinctly coarser than peripheral ones, 
u central and peripheral spinelets uniform. 
s one central spinelet both longer and stouter on most paxillae. 

S one central spine (1-5 mm long) abruptly enlarged on some scattered (mostly lateral) 

5. Shape of superomarginal paxillae: 

+ elongate. 

rounded or squarish. 

6. Alignment of inferomarginal plates: 

1 lateral. 

vl ventro-lateral. 

7. Inferomarginal spines: 

+ alternating on consecutive plates. 
all on the same levels. 

8. Number of large erect inferomarginal spines. 

9. Number of adambulacral spines. 

10. Number of valves on actinal pedicellariae (when present, - = absent). 

1 1 . Oral furrow pedicellariae: 

+ present. 

spine in the adult, this is probably evident proximally by R 12-15 mm and the other spines 
and spinelets grow and coarsen into their adult proportions at this size or soon after. Luidia 
sagamina aciculata Mortensen is exceptional in the development of the third adambulacral 
spine, which may be lacking altogether, especially in american specimens, judging from the 
few so far studied. Increase in numbers of spines also shows a growth gradient along the arm. 
In L. heterozona Fisher, for instance, only about four proximal adambulacral plates have the 
third spine enlarged at R 20 mm but there is even a fourth spine on the first 10-1 5 plates, and 
almost equal to the fully developed third one, by R c. 80 mm. 

Actinal and abactinal pedicellariae, when developed, probably first appear between R 1 5 
and 20 mm but oral furrow pedicellariae, for example in L. heterozona, are not fully 
modified from pairs of spines until R exceeds c. 50 mm. 


PREVIOUS WORKS. Useful relevant publications on the Luidiidae include the general 
monograph of Doderlein (1920), Nataf & Cherbonnier (1973) on the tropical West African 
area and Downey (1973) and Walenkamp (1976 and 1979) on the West Indian area. 

DISTRIBUTION. The species of Luidia are mostly restricted to the shelf, though several 
Atlantic taxa extend to the upper bathyal, including L. barbadensis to 430 metres, L. 
ciliaris to 400 metres, L. heterozona and L. sagamina aciculata to 975 metres and L. sarsi 
exceptionally to 1 300 metres. (The maximum depth for both L. heterozona and L. sagamina 
aciculata derives from Jean Charcot (Walda cruise) st. 40, c. 04 07' N, 01 35' W, off Ghana, 
975 metres, and is surprising since previous maxima for the two respectively were only 400 
and 150 metres, cited by Nataf & Cherbonnier (1973), the vast majority of records for both 
species being from 50-100 metres.) 

Geographically, five Atlantic taxa appear to be restricted to the american side of the 
ocean, namely L. barbadensis, L. clathrata, L. ludwigi scotti, L. patriae and L. senegalensis 
(the last despite its name), while L. atlantidea and L. ciliaris are entirely eastern (american 
records for L. atlantidea proving incorrect, see p. 168. The remainder are distributed as 


alternata N Carolina to N Argentina numidica Senegal to Zaire (Congo) 

barimae Venezuela to French Guiana heterozona Mauritania to Angola 

sagamina aciculataC?) N Carolina to sagamina aciculata Mauritania to 

Florida Strait Zaire & St Helena 

elegans E U.S.A. & Gulf of Mexico sarsi Norway to Mauritania & Azores 

eleganstf] S Brazil africana SW & S of southern Africa 

The relationships of these are discussed below to assess their respective ranks. 

LUIDIA Fofbes 

Luidia Forbes, 1839:123; Sladen, 1889:244-248; Fisher, 1911:105; Sussbach & Breckner, 

191 1 : 198; Doderlein, 1920 : 193-246; Bernasconi, 1943 : 3-5; A. M. Clark, 1953 : 379-380; Fell, 

1963 : 433; Downey, 1973 : 21-22; Tommasi, 1974 : 7; A. M. Clark & Courtman-Stock, 1976 : 43. 

Type species: Luidia fragilissi ma Forbes, 1839 (a synonym of Asterias ciliaris Philippi, 1837), by 

Hemicnemis Miiller & Troschel, 1840:105; Doderlein, 1920:217 (as subgenus). Type species: 

Asterias ciliaris Philippi, 1837. 
Petalaster Gray, 1840: 183: Doderlein, 1920:215 (as subgenus). Type species: P. hardwicki Gray, 

1 840, cited by Fisher, 1911 (though with Petalaster in synonymy), non P. Columbia Gray, designated 

by Fell, 1963. 

Luydia (lapsus) Diiben & Koren, 1846 : 254. 
Astrella Perrier in Milne-Edwards, 1882 :21. Type species: A. simplex Perrier, 1882 (a synonym of 

Luidia sarsi Diiben & Koren, 1 845), by monotypy. 
Senegaster, Maculaster, Alternaster, Armaster, Integraster, Denudaster, Penangaster & Quinaster- 

(subgenera) Doderlein, 1920:215-217, raised to generic rank by Fell, 433-434 (except for 


Luidia alternata (Say) 
Figs Ib, 2d, h, 3a-i,4, 5 

Asterias alternata Say, 1825 : 144-145. 

Luidia alternata: Liitken, 1859:42^13; Perrier, 1875:334-336 [1876:254-256]; Sladen, 1889: 
250-251, 740; Verrill, 1915 : 201-203 [non L. alternata bicolor}; Doderlein, 1920 : 241, 267-268, 
figs 7, 1 1; H. L. Clark, 1933 : 20, pi. 1 ; Bernasconi, 1943 : 14-15, pi. 3, figs 1,6, pi. 4, fig. 1; Engel & 
Schroevers, 1960 : 6; Ummels, 1963 : 97-98, pis 8, 10; Gray, Downey & Cerame-Vivas, 1968 : 138, 
fig. 6; Tommasi, 1970 : 8, pi. 8, fig. 24; Downey, 1973 : 23-24, pi. 2, figs A, B; Blake, 1973 : 33-34, 
pi. 6, figs 25-42; Walenkamp, 1976 : 29-32, figs 6, 7, pi. 3, fig. 3, pi. 4, fig. 4. 

Luidia granulosa Perrier, 1869 : 109-1 10, pi. 2, fig. 18. 

Luidia variegata Perrier, 1875:337[1876:257]. 

164 A.M.CLARK 

Luidia numidica Koehler, 1911 : 3, pi. 1, figs 8-11; 1914 : 167, pi. 4, fig. 7; Doderlein, 1920 : 235, 242; 

Nataf & Cherbonnier, 1 973 : 7 1-74, pi. 1 , figs A, C, pi. 5, figs 1 , 2, pi. 7, fig. 6, pi. 9, figs A-D. 
Luidia quequenensis Bernasconi, 1942 : 253; 1943 : 15-18, pi. 2, fig. 4, pi. 3, figs 2, 3, 7, pi. 5, figs 1, 2; 

1960 : 21-22, pi. 3, fig. 1 ; Carrera-Rodriguez &Tommasi, 1977 : 65. 
Luidia bernasconiae A. H. Clark, 1945 : 19-21; Gray, Downey & Cerame-Vivas, 1968 [?pt] : 138, fig. 

7; Downey, 1973:25. 

Luidia alternata var. numidica: Madsen, 1950 : 206-209, fig. 9. 
Luidia alternata numidica: A. M. Clark, 1953 : 388-389, pi. 41 , fig. 1 ; 1955 : 22; Engel & Croes, 1960 : 

11-1 2, pi. 6, figs. 1,2. 

R up to 175 mm. R/r ranging from c. 5-6/1 at R 3(MO mm to 7-9/1 at R> 70 mm. 

A species of Luidia with normally 5 arms; abactinal paxillae with about 4 longitudinal 
series each side also forming transverse rows with the matching but distinctly smaller 
superomarginal paxillae, though irregularities may occur where the paxillae of the second 
and third (sometimes other series) are markedly enlarged and bear a single stout central spine 
fringed by usually several rings of spinelets, the large spines similar in magnitude to the 
ambital inferomarginal spines, rarely some of the lateralmost series and many of the median 
paxillae may have a smaller central spine developed; inferomarginal plates primarily ventral 
in position, only narrowly visible dorsally, armed at the ambitus with 2 or 3 large spines, 
usually projecting approximately horizontally, and on the ventral side with 2-4 smaller and 
somewhat flattened spines among the spinelets: adambulacral plates with 3 large spines in a 
row at right angles to the furrow, sometimes a more or less enlarged spine proximal to the 
lateralmost one; elongate, 3- or 4-valved pedicellariae, present on some proximal actinal 
plates (rarely also on the outer ends of the first few adambulacral plates) the slender valves 
usually only meeting at the tips; shorter, more often bivalved pedicellariae sometimes 
present (especially in west african specimens) on some abactinal and/or superomarginal 
paxillae, no pedicellariae on the furrow face of the oral plates. Colour boldly patterned on 
the upper side with a dark pentagon on the disc and three to five transverse dark bands, 
brown, black, greenish or purple, on each arm, the remainder being yellow, cream or white. 

STATUS OF L. numidica. Madsen (1950) treated Luidia numidica Koehler as a west african 
variety of L. alternata and noted that the only difference appeared to be the smaller number 
of spine-bearing paxillae in numidica. A. M. Clark (1953) used subspecific rank for 
numidica but noted that a specimen from the Cape Verde Islands with relatively numerous 
spino-paxillae is very similar to some west Atlantic specimens of L. alternata. In 1973, Nataf 
& Cherbonnier treated numidica as a full species but without comparisons or any comment 
on its status. 

Koehler (1911) had only two specimens from West Africa, a poor dry one with R 52 mm 
and the broken holotype, r 13mm so R probably 80-100 mm. He remarked on the 
closeness to L. alternata but distinguished that species by the more numerous inferomarginal 
and paxillar spines, the more attenuated pedicellariae and the 'quite different' paxillae with 
the peripheral spinelets thin and elongated, not short, thick and granuliform. Study of c. 30 
and 15 specimens from west and east Atlantic respectively, mostly from the Gerda and 
Pillsbury collections, now indicates that, although the armament of the paxillae is somewhat 
variable in both east and west Atlantic specimens and is also subject to some extent to growth 
changes, there is a consistent difference in the armament of the smaller paxillae if total size is 
taken into account and also a significant difference in the number (often also the shape) of 
the large paxillar spines. Already in west african specimens at R 20-25 mm, over half the 
median paxillae have more than one (usually 2-4) short coarse similar central spinelets; at R 
40-50 mm these number 3-6 and at R c. 60 mm usually 4-8 (see Fig. 3h, i). In L. alternata 
from the western Atlantic, even at R 100 mm or more, many central paxillae still only have a 
single more or less coarser central spinelet, the maximum number being about 4. However, 
these large specimens may have some median paxillae with the peripheral spinelets arranged 
in two concentric rings of 6-10 each, the outer ones being more slender and cylindrical than 
the inner ring, the spinelets of which are transitional in shape to the central spinelet (or 
spine). In general, the shape of the peripheral spinelets can usually be called subclavate but 




Fig. 3 (a, b, d-p) Abactinal paxillae, (c, q-v) pedicellariae. (a-g) Luidia alternata alternata, (a) 
type material of L. quequenensis, Quequen, N Argentina, R 72 mm, median paxillae; (b) 
holotype of L. bernasconiae, R 100 mm; (c) the same, superomarginal pedicellaria; (d) Gerda st. 
1038, R 40 mm, median paxillae; (e) Oregon st. 42 1 5, R c. 60 mm, median paxillae; (f) Pillsbury 
st. 654, R 100+ mm; (g) Pillsbury st. 767, R c. 90 mm, superomarginal paxilla; (h, i) L. alternata 
numidica: (h) Pillsbury st. 248, R only c. 22 mm; (i) Cape Verde Is, R 55-60 mm; 0) L. 
heterozona heterozona, Pillsbury st. 68, Re. 100 mm, disc paxillae; (k) L. ciliaris, Naples, R 
140mm, inner lateral paxilla (above) and two median paxillae; (1) L. sagamina aciculata, 
Atlantide st. 61, R c. 90 mm, superomarginal paxilla (left) and lateral paxillae from above and 
the side (below); (m) L. sarsi sarsi, W. of Scotland, Re. 100 mm, median paxillae; (n) L. sarsi 
sarsi, paratype of L. africana, Porcupine st. 36, R c. 85 mm, disc paxillae; (o) L. sarsi elegans, 
Gerda st 456, R c. 60 mm, median paxillae; (p) L. atlantidea, Pillsbury st. 26, Re. 100 mm, 
lateral paxillae; (q) L. alternata alternata, Oregon st. 4190, actinal pedicellariae; (r) L. 
heterozona heterozona, Pillsbury st. 259 (left) and 49 (right), actinal pedicellariae in two views; 
(s) L. sagamina aciculata, Atlantide st. 6 1 , two actinal plates, one armed only by the pedicellaria, 
pedicellariae slightly foreshortened; (t) L. ludwigi scotti, paratype, actinal paxillae with 2-, 3- 
and 4-valved pedicellariae, foreshortened; (u) L. ludwigi scotti, Pillsbury sts 1384 & 842, lateral 
views of 4- and 3-valved actinal pedicellariae; (v) L. sarsi sarsi, Shetland Is, lateral views of 
actinal pedicellariae. 

166 A.M.CLARK 

alternate ones may be hardly at all expanded at the tips on the slightly larger paxillae. As for 
the large paxillar spines on the second and third lateral series of paxillae particularly, even in 
the Cape Verde Islands specimen mentioned in 1953 only c. 35%, at most, of these lateral 
paxillae have the large spines. In other specimens from West Africa the percentage is usually 
c. 20. The shape of the spines is also relatively squat, their length most often 1-2 mm, the 
longest (in Koehler's large holotype) 3 mm, the length/basal breadth usually less than 3/1. 
In american specimens such squat spines may also occur but usually the length/breadth 
ratio is4-(M'5/l and the length may exceed 3 mm even at R only 60 mm. 

The conclusion reached from these observations is that the specimens from the two sides 
of the Atlantic are conspecific but the west african form can be recognized as a distinct 
subspecies numidica of L. alternata, distinguished by the more 'plushy' superficial 
appearance of the abactinal paxillae, interspersed with shorter and fewer large spines. 

SYNONYMY. Luidia granulosa Perrier, 1869 was recognized as a synonym of L. alternata by 
Perrier himself in 1875 and L. variegata Perrier, 1875 by Verrill, 1915. In 1950, Madsen 
(cited by A. M. C., 1953 : 380) also referred L. bernasconiae A. H. Clark, 1945, to the 
synonymy but this was refuted by Downey in 1968 & 1973, who maintained that it has only 
two adambulacral spines, not three or four, and the non-spine-bearing paxillae have 5-10 
clavate peripheral spinelets compared with 10-16 slender acute ones in alternata, though on 
p. 23 of the 1973 paper she describes the peripheral paxillar armament of alternata as 
subclavate. The dried holotype of L. bernasconiae certainly shows swollen-tipped peripheral 
paxillar spinelets more clavate than usual for L. alternata but it regularly has three large and 
a fourth smaller adambulacral spine, as described by A. H. Clark, the common number for L. 
alternata. In view of the variability in shape of the paxillar armament in L. alternata already 
mentioned, it is difficult to maintain bernasconiae as distinct. If there are specimens from 
North Carolina as described by Gray, Downey & Cerame- Vivas (1968) with sporadic large 
spines but only two adambulacral spines, then these may have to be nomenclatorially 
distinguished but not as L. bernasconiae. The superficial difference in proportions between 
L. alternata and L. bernasconiae suggested by the photographs in the 1968 paper is 
correlated with the much larger size (R c. 135 mm, judging from the scale) of the specimen 
named alternata, and exaggerated by the greater reduction so that both appear of similar size 
though R is only c. 77 mm in the specimen named L. bernasconiae. 

With regard to Luidia quequenensis (and other species of Luidia described by her) 
Bernasconi, 1942 confusingly modified the terminology of the lateral abactinal paxillae used 
by Doderlein (1920, see pi. 18, caption) as follows: his 'Randpaxillen' (i.e. the supero- 
marginal paxillae) become her 'paxilas marginales'; his 'untere Seitenpaxillen' (the two 
[primary and secondary] outermost series of lateral paxillae) become her 'paxilas infero- 
marginales'; his 'obere Seitenpaxillen' (the third and fourth series or two outer series of 
median paxillae) become her 'paxilas superomarginales'. This odd terminology has unfor- 
tunately been perpetuated by Carrera-Rodriguez & Tommasi (1977). Bernasconi 
distinguished L. quequenensis from L. alternata in 1943 (pp. 5, 17) by the large number and 
coarse shape of the spines of the 'paxilas superomarginales', which should be the third and 
fourth series of paxillae from the true superomarginal ones. However, a specimen from 
Quequen, kindly presented to the British Museum by Dr Bernasconi, shows numerous 
spinopaxillae but more of these are in the second from outermost lateral series (what I call 
the primary row), though the third series also has a considerable number. However, there are 
many inconspicuous spineless paxillae also in these series so that even the second only has c. 
64% of spinopaxillae out of well over 100 paxillae counted on several arms. The largest 
number of consecutive spinopaxillae found was only 9. Similar frequencies of over 60% of 
spinopaxillae in the second or third lateral series may occur throughout the range of L. 
alternata alternata, for instance in some of the Challenger specimens from Bahia, NE Brazil, 
others from French Guiana (Oregon and Pillsbury collections) and from Georgia (the 
holotype of L. bernasconiae). 

NEOTYPE. In the event of future disagreement with the synonymies now proposed and in the 






T* ' 

Fig. 4 Luidia alternata alternata (Say), neotype, BM(NH) reg. no. 1937.5.9.6, Dry Tortugas. 

Dorsal view, xO'6. 

168 A.M.CLARK 

absence of Say's type material (apparently long-lost), it is very desirable that a neotype from 
the vicinity of Florida, the type locality, should be nominated. Accordingly I propose a 
beautiful specimen from the Dry Tortugas now in the British Museum collections, registered 
number 1937.5.9.6, collected by Colman and Tandy, shown in Figs 4 and 5. 

The arms vary in length so that R ranges from 150-1 75 mm; r is c. 19mm and 
R/r = 7 - 9-9'2/l. Spinopaxillae are fairly numerous on the proximal primary (second) lateral 
series but become more sporadic distally (c. 55% of 100+ primary paxillae from proximal 
parts of several arms are spine-bearing). The longest spines measure 4*0 mm in length. 
Progressively fewer of the third, fourth and fifth series of paxillae are also enlarged with 
spines but none of the first series or the adjoining superomarginal paxillae and very few 
midradial paxillae. The median paxillae without enlarged spines have at least one central 
and about 9 peripheral spinelets, the latter almost cylindrical and blunt-tipped but the 
slightly larger paxillae have up to 4 central and c. 20 peripheral spinelets, the latter in a 
double ring of which the inner ones are subclavate and transitional in shape to the shorter, 
coarser central spinelets. The inferomarginal plates mostly bear two large ambital spines up 
to 7 mm long, slightly irregular in position on consecutive plates but not regularly alter- 
nating. There are 2-4, usually 3, more irregular smaller spines on the ventral side of each 
plate. Most of the primary actinal plates bear a long slender 3- or sometimes 4-valved 
pedicellaria and proximally also one or two additional pedicellariae based on super- 
numerary platelets superimposed on the lateral ends of the adambulacral plates. Most 
adambulacral plates have 3 large spines and a smaller fourth one proximal to the third 
(lateral-most). On the proximal plates this fourth spine may reach a similar length to the 

DISTRIBUTION. L. alternata alternata from North Carolina to northern Argentina (c. 38 S); 
0-160 metres. 
L. alternata numidica from Senegal to Zaire (Congo), West Africa; 10-100 metres. 

Luidia atlantidea Madsen 
Fig. 3p 

Luidia africana: Koehler, 1911:19; Doderlein, 1920 : 288-289; Mortensen, 1925 (pt) : 178. [Non L. 

africana Sladen, 1889.] 
Luidia atlantidea Madsen, 1950 : 192-198, fig. 5, pi. 16, figs 1,2; A. M. Clark, 1953 : 393, 394; 1955 : 

22, 32; Nataf& Cherbonnier, 1973 : 76-80, pi. 1, fig. B, pi. 2, figs A, B, pi. 7, figs 1-5, pi. 9, figs E, F. 

[Non L. atlantidea: Downey, 1973 : 25 = L, sagamina aciculata.] 
Luidia sarsi: Studer, 1884 : 43. [Non L. sarsi Diiben & Koren, 1845.] 

DISTRIBUTION. On re-examination, the specimen from North Carolina mentioned under the 
name of L. atlantidea by Downey (1973) proved to be L. sagamina aciculata. It has a white 
band along each side of the paxillar area on the arms but this is situated on the two regular 
rows of lateral paxillae, not on the superomarginal series as in L. atlantidea. It has not been 
possible to check the Halpern record cited by Downey but the likelihood is that this too 
would have been a similar misidentification. 

The species is recorded from Atlantic Morocco to Zaire, including the Cape Verde Islands, 
10-80 metres. 

Luidia barbadensis Perrier 
Fig. 2b 

Luidia barbadensis Perrier, 1881 : 29; 1884 : 267 [non pi. 10, figs 7, 8, probabaly = L. sarsi elegans]; 

Verrill, 1915 : 205-207, pi. 24, fig. 1; Doderlein, 1920 : 216; H. L. Clark, 1941 : 25; John & A. M. 

Clark, 1954:145; Cherbonnier, 1959:170, fig. 3B, E, fig. 4; Downey, 1973:24-25; 

Carrera-Rodriguez & Tommasi, 1977 : 67-7 1 , figs 5-9. 
Luidia convexiuscula Perrier, 1881 : 30; 1884 : 268, pi. 10, fig. 6; Verrill, 1915 : 207. 



Fig. 5 Luidia alternata alternata, neotype. Ventral view of proximal part, x 1 -7. 

170 A.M.CLARK 

REMARKS. Records of five-armed specimens of L. barbadensis are probably based on 
specimens of L. sarsi elegans, which is partially sympatric with it. Certainly the five-armed 
specimen captioned as L. barbadensis by Perrier, 1884, pi. 10, figs 7, 8, appears to be 
elegans, which also differs in the much finer paxillae and the long marginal spines. All 36 
Gerda, Pillsbury and Oregon specimens of barbadensis which I have seen have 6 arms. 

DISTRIBUTION. Southern Florida and the immediately adjacent Gulf of Mexico, southern 
Bahamas, to SW Caribbean off Nicaragua and east and south to southern Brazil (c. 30 S), 
73-430 metres. 

Luidia ciliaris (Philippi) 
Fig. 3k 

Asterias ciliaris Philippi, 1837 : 144. 

Luidia fragilissima Forbes, 1839 : 123; 1841 (pt) : 135-140, fig. on p. 135 [7-armed specimens]. 

Hemicnemis ciliaris: Miiller&Troschel, 1840 : 104. 

Luidia ciliaris: Perrier, 1875 : 342 [1876 : 262]; Sladen, 1889 : 254; Ludwig, 1897 : 61-85, pi. 4, figs 1, 
2, pi. 6, figs 25-36; Sussbach & Breckner, 1911 : 209-210; Doderlein, 1920:287, figs 8, 17, 34; 
Koehler, 1921 : 55, fig. 41; Mortensen, 1927 : 90, fig. 89a; Madsen, 1950 : 205-206, fig. 8; Ursin, 
1960 : 30-3 1 ; Tortonese, 1965 : 148-150, fig. 68; Blake, 1973 : 40, pi. 10, figs 1-26. 

DISTRIBUTION. NE Atlantic, from southern Norway, the Skagerrak, Shetlands, Faeroe 
Channel, south to the Canary Islands and Azores (?Cape Verde Islands); 1-400 metres. 

Luidia clathrata (Say) 
Fig. la 

Asterias clathrata Say, 1825 : 142. [Non A. clathrata Pennant, 1777.] 

Luidia clathrata: Liitken, 1859:37-39; Perrier, 1875:332-333 [1876:252]; Sladen, 1889:253; 
Verrill, 1915 : 200-201, pi. 24, fig. 2; Doderlein, 1920 : 238, 239, 251-252, figs 1, 21; H. L. Clark, 
1933 : 19-20; Bernasconi, 1943 : 6-7, pi. 2, fig. 1; A. H. Clark, 1954 : 375; Ummels, 1963 : 95-97, 
pis 10, 11; Gray, Downey & Cerame-Vivas, 1968 : 139, fig. 8; Downey, 1973 : 22-23, pi. IJigsC, D; 
Blake, 1973:30-31, pi. 3, figs 1-28; Walenkamp, 1976:19-25, figs 2^t, 19a, pi. 3, fig. 2; 
1979: 11-12. 

NOMENCLATURE. Although A. H. Clark (1954) pointed out that Asterias clathrata Say was 
preoccupied by Pennant's earlier senior homonym, he commented that 'nothing would be 
gained by displacing this well established name' and so continued to use it for this common 
West Indian species. Unfortunately, under the rules of nomenclature (Article 59(a)) such a 
junior primary homonym 'must be permanently rejected', that is unless the International 
Commission can be prevailed upon to use its plenary powers to suppress the earlier name 
(itself a synonym of Asterias rubens Linnaeus, 1758), which action is being requested in a 
separate proposal. 

DISTRIBUTION. North Carolina (??New Jersey), Bermuda and the Gulf of Mexico to southern 
Brazil (c. 3 1 S); 0-175 metres but rarely below 70 metres. 

Luidia heterozona Fisher 
Figs2f,i,j,3j, r 

Luidia heterozona Fisher, 1940:265-268, fig. M4, pi. 23; Madsen, 1950:203-205; John & A. M. 

Clark, 1954 : 148; A. M. Clark, 1955 : 33; Cherbonnier, 1963 : 182; Nataf & Cherbonnier, 1973 : 

74-76, pi. 4, figs A, B, pi. 5, figs 3-6, pi. 9, figs I, J; Sibuet, 1975 : 284, 288. 
Luidia mortensem Cadenat, 1941 : 53-67, figs 1-3; Cherbonnier, 1963 : 182. 
Luidia barimae John & A. M. Clark, 1954: 145-148, figs 3, 4, pi. 6, fig. 2; Downey, 1973 : 25; 

Walenkamp, 1976: 18-1 9, pi. 1, figs 1-3. 

R up to 255 mm; R/r 7-1 1/1. Of over 150 specimens from both sides of the Atlantic taken by 
the Gerda, Pillsbury and Oregon, three have 9 arms, all the rest 10, though Madsen (1950) 
found 2 of the 37 Atlantide specimens to have 9 arms and another only 8. 


A species of Luidia with usually 10 arms, relatively slender and attenuated; abactinal 
paxillae all relatively small and rounded but rather variable in size medially, some slightly 
larger ones intermingled; laterally two matching longitudinal series each side, outnumbering 
the adjacent more elongate superomarginal paxillae by 15-18/10; inferomarginal plates 
mainly lateral in alignment, bearing up to four spines in L. heterozona barimae but rarely 
more than three in L. heterozona heterozona, the spines erect, long and acute, the uppermost 
(at least on alternate plates) projecting vertically upwards and the uppermost or the second 
the longest, the spines alternating fairly regularly in level on successive plates and often also 
in number 4/3 or 3/2, the plates unusually bare of spinelets between the large spines except 
at the edges; actinal plates unusually elongate, partially overlying the abradial ends of the 
adambulacrals; adambulacral plates mostly with three spines in line at right angles to the 
furrow but in larger specimens, R>70 mm, c. 10-15 proximal plates may have a fourth 
spine enlarged proximal to the abradial one; pedicellariae bivalved throughout, often very 
abundant, especially on the paxillae (Fig. 3j) where they are relatively short and rounded, 
their edges almost fully contiguous, actinal pedicellariae longer, sometimes broadened at the 
tips, numbering up to three on some proximal actinal plates, similar ones on the infero- 
marginals and oral plates while one to three more spiniform pedicellariae also occur on the 
furrow face of each oral plate near the mouth. Colour in life dark chocolate brown above 
except for a white band along the upper edge of each arm (possibly the superomarginals), the 
inferomarginal spines white with brown bases. 

STATUS OF L. barimae. Luidia barimae was established on the basis of two poorly preserved 
specimens from off Venezuela. Walenkamp (1976) has recorded nine others from Surinam, 
evidently also in poor condition. About 50 further specimens from the Pillsbury collections 
off the north coast of South America have now been studied, as well as more than 100 from 
West Africa of L. heterozona. Though many of these are also poor, some are good enough to 
show that there are two matching regular longitudinal series of lateral paxillae adjacent to 
the superomarginal paxillae in the american and well as african specimens. Also the supero- 
marginal paxillae are similarly markedly larger than the other paxillae (Fig. 2f). This leaves 
only the number of inferomarginal spines as a possible distinction between the two, L. 
barimae being described as having four spines proximally, then three, whereas no more than 
three, more often three and two, are said to be found in L. heterozona by Fisher, Madsen and 
Cadenat. However, one large Pillsbury specimen from the Gulf of Guinea, West Africa, with 
R at least 1 50 mm, was found to have four inferomarginal spines on a few proximal alternate 
plates, while conversely a specimen from Venezuela with Re. 120 mm has no more than 
three spines, though these reduce to two on the alternate distal plates. Certainly there is no 
justification for more than a subspecific distinction between specimens from the two sides of 
the tropical Atlantic and the minor difference in the marginal spines is clearly correlated 
with different growth rates. 

DISTRIBUTION. L. heterozona heterozona from Cap Blanc, Mauritania, south to Elephant 
Bay, Angola; 28-975 metres. As noted above (p. 163), apart from Sibuet's Walda Expedition 
record (1975), the greatest depth recorded is 400 metres. 

L. heterozona barimae from off the Gulf of Maracaibo, western Venezuela (c. 
1234'N:71 10'W) eastwards to French Guiana (0607' N : 5219' W); 38-90 (7100) 

Luidia ludwigi scotti Bell 

Figs2a, e, 3t, u 

Luidia scotti Bell, 1917 : 8-9; A. M. Clark, 1953 : 383-385, fig. 3, pi. 40, fig. 1; John & A. M. Clark, 

1954 : 144; Carrera-Rodriguez & Tommasi, 1977 : 62, 65-66. 

Luidia doello-juradoi Bernasconi, 1941 : 117; 1943 : 8-11, pi. 1, fig. 3, pi. 2, figs 2, 3, pi. 3, figs 4, 5. 
Luidia rosaurae John & A. M. Clark, 1954: 142-145, figs 1,2, pi. 6, fig. l;Jangoux, 1978 : 95. 
Luidia ludwigi: Walenkamp, 1976 : 32-37, fig 9, pi. 2, figs 1, 2, 4, pi. 4, fig. 3. 

R up to 90 mm; R/r 5'3-8'l/l, mean of 9 specimens 7*0/1 . 

172 A.M.CLARK 

A species of Luidia with normally 5 arms: abactinal paxillae with two longitudinal 
rectangular or squarish lateral series each side, matching each other and also the adjacent 
slightly smaller superomarginal paxillae, central paxillar spinelets distinctly coarser than the 
peripheral ones and in smaller specimens, R<c. 40 mm, where the median paxillae often 
have only a single central spinelet, this may stand out from the surface slightly but not as 
much as in L. sagamina; larger specimens with multiple central spinelets have them 
relatively shorter and making a uniform surface; inferomarginal plates mainly ventral in 
alignment, bearing one or two (in the largest specimens, R c. 90 mm, sometimes three) stout 
ambital spines, the upper one more or less shorter than the lower when there are two, on the 
ventral side 4 or 5 much smaller, almost cylindrical, but slightly tapering spines; 
adambulacral plates with 4 large spines, the abradial two in line parallel to the furrow; 
pedicellariae 3- or 4-valved on at least the more proximal actinal plates, the valves varying in 
shape from broadly spatulate to evenly tapering and triangular (see Fig. 3t, u), sometimes 
bivalved pedicellariae present on the paxillae and inferomarginal plates, absent from the 
furrow face of the oral plates. Colour (recently dried) patterned above with dark rose- 
coloured patches on the centre of the disc and bands across the arms, sometimes linked by 
continuous colour midradially. 

SYNONYMY. The poorly-preserved holotype and two paratypes of Luidia rosaurae John & 
Clark, 1954, from off Venezuela, were thought to differ from L. scotti Bell, 1917, of southern 
Brazil in having only a single large ambital inferomarginal spine, finer ventral armament and 
more numerous pedicellariae. Walenkamp's material from Surinam (1976) and a number of 
Pillsbury and Gerda specimens from northern South America and the Florida Strait now 
studied, show that there is considerable variation in the development of the second (upper) 
inferomarginal spine, the thickness of the ventral spines and the frequency of the 
pedicellariae; consequently L. rosaurae cannot be maintained as specifically distinct. 
Walenkamp deferred a decision on this point in default of material from the type locality of 
L. scotti. Simultaneously, he rejected the synonymizing of Luidia doellojuradoi Bernasconi, 
1941 with L. scotti, proposed by A. M. Clark (1953), on the grounds that its marginal 
paxillae are larger than the lateral ones and the R/r ratio is only 7/1 . However, he was misled 
in this by Bernasconi's peculiar terminology of the lateral and superomarginal paxillae (see 
p. 166), the true superomarginal paxillae of L. doellojuradoi being in fact smaller than the 
lateral ones. Also in the 1 5 Terra Nova specimens of L. scotti the R/r ratio ranges upwards 
from 7/1; in the lectotype (incorrectly called holotype in 1953) it is 7*5/1. Thanks to Dr 
Bernasconi, there are two specimens which she named as L. doellojuradoi in the British 
Museum collections. Apart from the greater extent of the actinal pedicellariae on the arms 
and the occurrence of some inferomarginal pedicellariae characters not considered to be of 
specific weight no significant differences from L. scotti are apparent. The ventral 
armament of the inferomarginal plates has become appressed during preservation, which 
lends a rather coarser appearance than is shown by the well preserved type material of L. 

It is also possible that L. patriae Bernasconi will prove to be synonymous with scotti. 

Walenkamp (1976) percipiently noted the close affinity between L. ludwigi Fisher, 1906, 
from California and specimens from the north coast of South America conspecific with L. 
rosaurae, referring the latter to the synonymy of L. ludwigi. Fisher's holotype has R 
1 10-1 1 5 mm and superficially resembles the larger Atlantic specimens seen except that it is 
rather coarser. A paratype of L. ludwigi with R 72 mm compares closely with a paralecto- 
type of L. scotti of almost the same size but there appears to be a significant difference in the 
number of paxillar spinelets, the median paxillae of ludwigi rarely having less than 6 coarse 
ones whereas in L. scotti the number is most often only about 3. Possibly there is also a 
colour difference, Fisher having described L. ludwigi (at least when freshly preserved) as 
being reddish above 'sometimes mottled with lighter', whereas Atlantic specimens seem to 
be more boldly patterned dark (?red) and light. In conjunction with the geographical 
difference, these small differences are now considered to warrant a subspecific distinction. 


One further nomenclatural threat still remains. Doderlein (1920), in listing L. ludwigi, 
noted that it is possibly (vielleicht) a synonym of L. armata Ludwig, 1905, from the Gulf of 
Panama. Accordingly, one of the two intact syntypes of L. armata from the U.S. National 
Museum has been studied; it has R 47 mm and is slightly decalcified and flaccid. The 
armament of the paxillae is more or less appressed, emphasizing the spiniform shape of the 
central spinelets. Although the larger Pacific specimens of L. ludwigi have markedly coarser 
and shorter central spinelets, tending to appear granuliform, smaller Atlantic specimens 
from the vicinity of Florida with R c. 40 mm, have a very similar relatively elongate 
armament and also agree in the shape and arrangement of the paxillae, plates and spines. 
However, the syntype of L. armata differs in having a very large, highly modified bivalved 
pedicellaria on each oral plate a feature somehow overlooked by Ludwig and not found in 
the type material of L. ludwigi seen or any Atlantic specimens. It also has relatively small 
actinal interradial areas with no sign of a second row of actinal plates. At R 40 mm, L. scotti 
already has a single median interradial actinal plate distal to the main series and in larger 
specimens, R 50-60 mm, there are one or two such plates each side of the interradius. These 
two characters, but particularly the first, indicate that there are two distinct species in the 
East Pacific. 

DISTRIBUTION. Eastern Gulf of Mexico west of Florida, (c. 21 { N : 84 W) and Florida Strait, 
also the northern and eastern coasts of South America from Venezuela to northern 
Argentina (c. 39 S); 33-126 (7135) metres. The northernmost record from 85 miles west of 
St. Petersburg, Florida, is thanks to Dr. K. Serafy and with seven Gerda stations in the Florida 
Strait provides a considerable extension of the range from South America. About 35 
specimens from 12 Pillsbury stations came from Venezuela to French Guiana and also 
served to extend the bathymetric range both up and down, the three shallowest 33, 36 and 
42 metres being from Surinam and French Guiana. There is also a specimen labelled as 
being from Pillsbury st. 1384, which is in the Puerto Rico Trench (c. 1945' N : 67W W) at 
7956 metres! Although it has relatively long arms (R/r 65/8 mm = 8'l/l), about the 
maximum found in scotti, the paxillar structure, form of the pedicellariae, general armament 
and banded colour pattern leave no doubt of the identification, but the locality is evidently a 

Ludiapatriae Bernasconi 

Luidia patriae Bernasconi, 1941 : 1 17-1 18; 1943 : 11-13, pi. 1, figs 1,2. 

R 72 mm (holotype); R/r 7-2/1 . 

A species of Luidia with normally 5 arms; abactinal paxillae with the two lateralmost 
series forming transverse rows with the matching superomarginal paxillae and all rectangular 
or squarish in shape: inferomarginal plates mainly ventral in alignment, bearing two slightly 
flattened and slightly curved large spines near their upper ends at the ambitus, the lower one 
larger, and on the ventral side about six pointed spines, cylindrical or somewhat flattened; 
adambulacral plates with four large spines, the two abradial ones in line parallel to the 
furrow: pedicellariae absent throughout. Colour (dried) dark rose above with darker areas in 
the centre of the disc and on the median parts of the arms. 

AFFINITIES. L. patriae is very similar to L. ludwigi scotti, of which Bernasconi had as many 
as 80 specimens (as L. doello-juradoi), though only six or seven of L. patriae, which she 
supposed to differ in the absence of pedicellariae (not now thought to be a character of 
specific weight), the form, number and arrangement of the ventral inferomarginal spines and 
in lacking the two or three enlarged spinelets abradial to the four main adambulacral spines 
described for L. doello-juradoi. Possibly L. patriae will also prove to be a synonym of L. 
ludwigi scotti. 

DISTRIBUTION. Northern Argentina. 34-37^ S [?also from Uruguay, 33S]; 100-126 metres. 

174 A.M.CLARK 

Luidia sagamina aciculata Mortensen 
Fig. Id, 2k, 31, s 

?Luidia alternate, bicolor Verrill, 1 9 1 5 : 203 , pi. 1 2, fig. 1 . 

Luidia sagamina: Downey, 1973 : 24. 

Luidia aciculata Mortensen, 19336 : 425-426, fig 7, pi. 20, figs 7-12; Fisher, 1940 : 268-269, fig. M5; 

Nataf & Cherbonnier, 1973 : 80-82, pi. 3, figs A, B, pi. 6, figs 1-6, pi. 9, figs G, H: Sibuet, 1975 : 284, 


Luidia sagamina var. aciculata: Madsen, 1950 : 199-203, figs 6, 7. 

Luidia sagamina aciculata: A. M.Clark, 1955 : 33; A. M. Clark & Courtman-Stock, 1976 : 23,32,45. 
Luidia atlantidea: Downey, 1973 : 25 [Non L. atlantidea Madsen, 1950.] 

Rupto 140 mm; R/r 7-0-10-5/1. 

A species of Luidia with normally 5 arms; abactinal paxillae all with an enlarged median 
spinelet (sometimes replaced by a pedicellaria) both stouter and longer than the remaining 
spinelets; two (or in the largest specimens three) longitudinal series of lateral paxillae also 
matching transversely but outnumbering the much longer superomarginal paxillae by 
16-19/10; inferomarginal plates mainly lateral in alignment, bearing two or three, 
sometimes four large spines, tending to alternate in number and position on consecutive 
plates, the uppermost spine the longest and projecting vertically upwards in its natural 
position, at least on every second plate; adambulacral plates with two or three large spines in 
a line at right angles to the furrow, when only two, the abradial one is relatively large; 
pedicellariae usually present on at least the proximal actinal plates, where they are more or 
less broadened basally and tapering so that the individual valves are triangular, though not 
closely contiguous except terminally, abactinal pedicellariae bivalved and shorter, some- 
times absent but one or more elongate bivalved pedicellariae always present on the furrow 
face of each oral plate near the mouth, though not fully developed from spinelets in small 
specimens. Colour in life purple to purple-brown above, white below, marginal spines dark 
brown basally. 

SYNONYMY. Madsen (1950) decided that Luidia sagamina Doderlein, 1920 from Japan and 
L. aciculata Mortensen, 1933 from St Helena are conspecific, following comparison of 
Mortensen's type material with a specimen from Sagami Bay and others collected off West 
Africa by the Atlantide. Although he retained aciculata as a subspecies, he could not find any 
real difference between it and sagamina. This prompted Downey (1973) to drop aciculata 
and use L. sagamina for some american specimens, though conversely Nataf and 
Cherbonnier in the same year retained aciculata at the specific level, without explanation. 
Madsen had also referred to L. sagamina aciculata a specimen from Durban, Natal and in 
1976 (Clark & Courtman-Stock) I retained this terminology, though without having seen any 
specimens from either Japan or Natal. Since then, further echinoderm material from off 
Natal (A. M. Clark, 1977) has shown that the fauna of SE Africa has much in common with 
that of southern Japan and the East Indian area. This would imply that the Durban specimen 
should be named L. sagamina sagamina if two subspecies are to be recognized, as seems 
desirable to me in the absence of precedent from related species of moderate depths common 
to similar areas of the Atlantic and Indo-West Pacific. 

A further complication is raised by the possibility that Luidia sagamina aciculata is 
conspecific with the (two?) specimens from the Florida Strait which Verrill (1915) called 
Luidia alternata variety bicolor. VerrilFs drawings show relatively small single spines (or 
enlarged spinelets) on all the abactinal paxillae, quite different from the sporadic and much 
larger spines of L. alternata. Also the lateral paxillae outnumber the inferomarginal plates 
(the superomarginals were not distinguished by the artist) and the inferomarginal spines are 
regularly alternating on consecutive plates, as in L. sagamina and aciculata. Downey (1973) 
recorded a small specimen (R in fact 2 1 mm not 8 mm as given in her paper) from South 
Carolina as L. sagamina and mentioned another (p. 25) which she referred to L. atlantidea 
because of light lateral bands along the arms but which also proves to be conspecific with the 
first. A further specimen, R 24mm, from Gerda st. 1039 in the Florida Strait may be 


Halpern's supposed L. atlantidea, mentioned by Downey. Unfortunately the Bahama 
Expedition's specimens of L. alternata var. bicolor cannot now be found at the University of 
Iowa. Since the revival of bicolor as a specific name would mean that both L. sagamina 
sagamina and aciculata became junior synonyms and there is an element of doubt about the 
identity, it is desirable that the name bicolor should be formally set aside. 

Verrill's specimens evidently had only two adambulacral spines, which is also the case in 
the small Silver Bay and Gerda specimens. Mortensen's holotype of L. aciculata from St. 
Helena, Fisher's Discovery specimen from off the Congo (Zaire), a Pillsbury specimen from 
the Gulf of Guinea (Fig. 2k) and an Atlantide west african specimen all usually have three 
adambulacral spines. Unfortunately, Madsen recorded four other Atlantide specimens with 
only two such spines and this is also true of two other Pillsbury Guinea specimens. Clearly, 
this character of adambulacral armament cannot be used to distinguish yet another 
american, as opposed to african, subspecies, comparable to the subspecies of L. heterozona 
and L. alternata, which would make for consistency of taxonomic treatment. The present 
american specimens are too small and too few to suggest other possible differential 

DISTRIBUTION. North Carolina to south and west Florida and from Cap Blanc, Mauritania 
south to Zaire and from St. Helena, 20-975 metres. As noted above (p. 163), apart from 
Sibuet's Walda Expedition record (1975), the greatest depth recorded is 1 50 metres. 

Luidia sarsi Diiben & Koren 
Figs Ic, 2c, g, 3m-o, v, 6 

Asteriassp. aff. A. aranciaca: M. Sars, 1835 : 39. 

Luidia fragilissima (pt, five-armed specimens) Forbes, 1841 : 135-140. 

Luidia Sarsii Dttben & Koren, 1845:113; Perrier, 1875:342 [1876:262]; Sladen, 1889: 258; 
Sussbach & Breckner, 1911:210. 

Luydia Savignyi (pt) Diiben & Koren, 1 846 : 254, pi. 8, figs 23, 24. 

Luidia elegans Perrier, 1875 : 336-337 [1876 : 256]; 1884 : 269[?], pi. 10, fig. 7 [and probably also fig. 
8], captioned 'L. barbadensis'; Verrill, 1885 : 543, pi. 13, fig. 39; 1915 : 203-205, pi. 16, figs4,4a, pi. 
19, fig. 1; Doderlein, 1920:289-290, fig. 36; Gray, Downey & Cerame-Vivas, 1968 : 140, fig 9; 
Downey, 1973 : 25, pi. 3, figs C, D; Blake, 1973 : 40-41, pi. 10, figs 27-52; Carrera-Rodriguez & 
Tommasi, 1977 : 71-75, figs 10-12. 

Astrella simplex Perrier, 1882 : 21, fig. 25; 1894 : 193-194, pi. 14, fig. 3. 

Luidia africana Sladen, 1889 : 256-258, pi. 44, figs 1, 2, pi. 45, figs 1, 2; Koehler, 1923 : 132; H. L. 
Clark. 1923 : 252; 1925(pt) : 8; Mortensen, 1933<z : 239-240, figs 3, 4; Madsen, 1950(pt) : 188-192, 
fig. 4, pi. 16, figs 3, 4; A. M. Clark, 1952 : 195; 1953 : 393-394, figs 10, 1 1; Nataf & Cherbonnier, 
1973:79, pi. 8; A. M. Clark & Courtman-Stock, 1976:23, 30, 44, fig. 26. [Non L. africana: 
Koehler, 191 1 : 19; Doderlein, 1920 : 228, nee Mortensen, 1925 : 178 = L. atlantidea.} 

Luidia sarsi: Bell, 1893 : 72; Koehler, 1895 : 320, pi. 9. figs 6, 7; Ludwig, 1897 : 85-104, pi. 4, figs 2, 3, 
pi. 7, figs 1-12; Koehler, 1909 : 59; Doderlein, 1920 : 288-289, fig. 35; Koehler, 1921 : 57, fig.; 
Mortensen, 1925 : 178 [Isarsi ace. Madsen, 1950]; 1927 : 69, fig. 39b; Tattersall & Sheppard, 1934 : 
36-55 (larvae); Madsen, 1950: 187; Tortonese, 1965: 150-152, fig. 69. [Non L. sarsi: Studer, 
1 884, = L. africana ace. Doderlein, 1920 but much more likely = L. atlantidea.] 

Luidia paucispina von Marenzeller, 1 893 : 3-4, pi. 1 , fig. 1 . 

SIZE. The maximum size varies in different parts of the range. In NE Atlantic specimens (L. 
sarsi sarsi) R reaches only c. 1 10 mm (BM coll., W of Scotland) but the same subspecies from 
the Mediterranean, L. sarsi africana from South Africa and L. sarsi elegans from America 
may reach 180-190 mm R; R/r 5'0-10'5/1, usually 7-5-10-0/1 at R>90 mm. 

DIAGNOSIS. A species of Luidia with normally 5 arms, becoming long and evenly tapering in 
large specimens, R>90 mm, but somewhat petaloid in smaller ones; abactinal paxillae 
relatively small and rounded, with two matching slightly coarser longitudinal lateral series 
each side but these are inconspicuous, outnumbering the markedly elongate superomarginal 
paxillae by 17-20/10, paxillar armament of fine spinelets of fairly uniform length, 
median paxillae mostly with only a single central spinelet, sometimes distinctly coarser in L. 

176 A.M.CLARK 

sarsi sarsi where the peripheral spinelets are also less attenuated than in L. sarsi africana and 
L. sarsi elegans (see Fig. 3m, n, o); inferomarginal plates mainly lateral in alignment, bearing 
2-4 usually 3, large pointed spines, showing some tendency to alternate in position and 
number on successive plates, the two upper spines similar in length, the uppermost often 
longer when not at its highest position; actinal plates distinctly broadened and keeled trans- 
verse to the arm axis; adambulacral plates with three large spines in series transverse to the 
furrow and sometimes one (or two) enlarged spinelets proximal to the lateralmost; 
pedicellariae bivalved with fairly broad rounded tips, usually almost circular in cross 
section but the valves sometimes rather flattened towards the tips, common on the median 
abactinal paxillae of L. sarsi africana and L. sarsi elegans, occasional in L. sarsi sarsi but 
longer actinal pedicellariae usually present in all three subspecies proximally, with one on 
the furrow face of each oral plate (exceptionally represented by two hardly modified spines). 
Colour in life brownish yellow, reddish or orange above, pale below, arms medially and 
marginal plates darker, especially in L. sarsi africana. 

STATUS OF L. africana AND L. elegans. These two taxa of Sladen, 1889 and Perrier, 1875 are 
here regarded as conspecific with L. sarsi. 

Confusion in the limits between L. sarsi and L. africana has arisen because the type 
material of the latter included one specimen from Atlantic Morocco besides the main 
sample from South Africa, one of the latter conforming to the size and description given by 
Sladen, as I noted in 1953 (p. 393). The moroccan specimen has fairly numerous globose 
pedicellariae on the abactinal paxillae, as in the southern specimens. Although such 
pedicellariae were thought to be rare in north european specimens of L.. sarsi, some larger 
specimens from west of Scotland collected since 1953 were found to have more or less 
numerous pedicellariae (Fig. 3m), sometimes two or three on a single paxilla. It is significant 
that there are no reliable identifications of L. africana from any localities between Cape 
Verde and Luderitz Bay, Namibia. Despite a critical comparison by Nataf & Cherbonnier 
(1973) out of 213 five-armed specimens of the ci/zam-group (excluding L. sagamina 
aciculatd) from this area, every one was found to be referable to L. atlantidea and not to L. 
africana. Doderlein's record (1920) of L. africana from the Cape Verde Islands (based on two 
specimens named L. sarsi by Studer in 1884) is clearly in mistake for L. atlantidea since he 
particularly notes the relative coarseness of the central paxillar spinelets and the occurrence 
of pedicellariae on the superomarginal but not the abactinal paxillae the reverse of what is 
found in L. africana, where both central and peripheral paxillar spinelets are uniformly fine. 
Madsen (1950) has referred Mortensen's larger moroccan specimens also to L. atlantidea, 
leaving under the heading of L. africana only some smaller moroccan ones which 'might just 
as well belong to L. sarsi' and three fragmentary Atlantide specimens which 'are not 
immediately recognizable as L. africana'. Zoogeographically, it would be expected that the 
distribution of a northern species also found in the Mediterranean, could well extend south- 
wards to some extent in north-west Africa, as with Marthasterias glacialis, for instance, 
which also occurs in South Africa but is unrecorded between Cape Verde and Cape Town. 

There are in the British Museum collections six specimens from near Cap Blanc 
(Discovery stations 8005 and 8020, 101 and 261-297 metres) with R 32 to c. 140 mm. These 
are certainly not L. atlantidea, having uniformly fine paxillar armament, abactinal rather 
than superomarginal pedicellariae, no white lines emphasizing the positions of the supero- 
marginals but darker brown midlines to the arms, much as in L. sarsi and L. africana. These 
specimens are a little unusual in having the pair of lateral spines on the furrow face of each 
oral plate hardly modified, if at all, into a pedicellaria. One exception is the smallest one, 
surprisingly since it is usually in larger specimens of species such as L. heterozona that this 
pedicellaria is more modified. Nearly all the specimens of L. sarsi, africana and elegans seen 
have quite a well-developed oral furrow pedicellaria, though the two spines from which it is 
modified may be more or less unequal in size. A second feature of some of these Cap Blanc 
specimens is that some proximal superomarginal paxillae are unusually broad, almost 
square, rather than consistently elongated, squarish paxillae being a characteristic of L. 


atlantidea. This is naturally limited to the plates adjacent to those inferomarginals where the 
highest spine is not at the upper edge of the plate. 

In describing L. africana, Sladen (1889) cited seven supposed differences from L. sarsi, the 
larger size (R up to 160 mm in the type material), the more even taper of the arms, the 
smaller size of the median paxillae, the lesser prominence of the central median paxillar 
'granule' (spinelet), the greater length of the uppermost inferomarginal spine relative to the 
second spine, the greater length of the third (lateralmost) adambulacral spine, achieving 
equality with the second spine, and the less pointed form of the actinal pedicellariae. Subse- 
quently Madsen (1950) noted that abactinal pedicellariae are not only much more often 
found in L. africana than in sarsi, where they are rare, but also that their position, when 
present in sarsi, is peripheral, while those of africana are central on the paxillar tabulum, so 
that the pedicellariae of sarsi tend to be seen in profile rather than from the top. He also 
noted that the inferomarginal spines of L. africana differ in having dark pigmented skin 

As far as size is concerned, although in Norway, the type locality of L. sarsi, larger 
specimens usually have R only 50-60 mm, around Scotland it may reach c. 1 10 mm and in 
the Mediterranean reputedly c. 180 mm (diameter 35 cm). Such larger specimens also have 
the arms more attenuated and evenly tapering and there is greater disparity in the size of the 
additional median paxillae. As for the relative lengths of the inferomarginal and 
adambulacral spines, these are dependent to some extent on growth, the lateralmost 
adambulacral spine being retarded in growth relative to the second spine, while it would be 
more correct to say that the two upper inferomarginal spines of L. sarsi are similar in length, 
the uppermost one often slightly longer, especially on those plates where it is not in its 
highest position, as is the case in the lectotype now proposed (see below), even though R is 
less than 40 mm. At R 100+ mm the third spine approximates in length to the other two in 
specimens from all localities, a characteristic thought by Doderlein (1920) to be diagnostic of 
L. elegans. 

With regard to the occurrence of abactinal pedicellariae, in 1953 I noted that in four out of 
ten northern specimens of L. sarsi studied, some pedicellariae were present. As mentioned 
above, subsequent Scottish collecting has yielded material with pedicellariae as numerous as 
in the moroccan paratype of L. africana and some south african specimens. In 1953 I also 
contended that the pedicellariae are not really central on the paxillae of L. africana (or L. 
elegans) either but that this may appear so when the paxillae are crowded in preservation 
and the valves are coarser, pushing the pedicellariae into a more nearly vertical alignment on 
the tabulum. There is rarely a complete circlet of spinelets, as shown in Mortensen's fig. 3 
(1933a) and Madsen's fig. 4c, d (1950). The shape of the pedicellariae, both abactinal and 
actinal ones, varies to some extent in different specimens from both north and south. Viewed 
end-on, the outline is usually approximately circular and the shape appears globular but 
may be slightly oval in one plane or the other if the valves are either thickened or broadened. 
The very broad shapes of both kinds of pedicellaria shown in Madsen's fig. 4h, i and k (1950), 
the abactinal ones cockle-shaped and the actinal ones with the individual valves bat-shaped 
(fan-shaped according to Madsen) seems to be exceptional even among south african 
specimens, none of the ten now studied having such an exaggerated form. Usually their 
actinal pedicellariae are either tapered in both side views (like a bishop's mitre) or else the 
tips are blunter and often thicker, a range also shown by northern specimens. 

Finally, the shape of the abactinal paxillar spinelets needs to be compared. In many 
northern specimens of L. sarsi the majority of median paxillae do have the single central 
spinelet distinctly coarser than the peripheral ones, appearing almost granuliform as 
foreshortened. This is particularly, but not exclusively, true of smaller specimens, including 
the lectotype, but in other specimens the thickness of all the paxillar spinelets is similar. A 
comparable modification of the central paxillar spinelet is noted above in some smaller 
specimens of L. ludwigi scotti and of course in a much more exaggerated form in L. 
sagamina aciculata, where the central spinelet is more or less markedly elongated as well as 
being much thicker than the peripheral spinelets. In general, the paxillar spinelets of south 

178 A.M.CLARK 

african specimens are appreciably more attenuated than those of european specimens and 
none have been observed to show a coarsening of the central spinelet, though it should be 
emphasized that no specimens with R< 70 mm are available. 

To sum up, in comparison between specimens from South Africa and those from Europe 
and NW Africa, no differences of specific weight can be found. There are slight differences in 
the armament of the paxillae in some specimens, the southern ones tending to have a finer 
and more uniform armament but this seems to be shared by some moroccan specimens and is 
always very subtle. Although the frequency of pedicellariae seems to increase further south, 
only in occasional south african specimens with extremely broad pedicellariae are these 
organs distinctive. It is therefore doubtful whether the name africana is worth retaining for a 
subspecies from South Africa but the distribution gap in west Africa coupled with these 
minor morphological differences may justify it. 

With regard to Luidia elegans from the american side of the Atlantic, the only comparison 
given by Perrier was of the ventral side with that of L. alternata just described, the difference 
emphasized being the bivalved rather than trivalved pedicellariae. In Doderlein's key (1920), 
he distinguished L. elegans from the much more closely related L. sarsi and L. africana by 
the equal length of all three inferomarginal spines being supposedly limited to L. elegans, 
which was also said to be the only one to have numerous paxillar pedicellariae. This 
comparison was evidently made primarily on the evidence of a single small specimen of L. 
sarsi (R 29 mm), one of L. elegans with R 145 mm and two specimens from the Cape Verde 
Islands which he thought to be L. africana but are more likely L. atlantidea. Inevitably, in 
smaller specimens the lowest inferomarginal spine, if developed at all, is relatively smaller, 
while at most stages of growth the two upper spines are similar in length, as discussed above. 
At Re. 100 mm in European specimens all three spines may be about equal, while the 
occurrence of pedicellariae is clearly variable and of little taxonomic weight. 

However, there does appear to be a significant difference in the number of abactinal 
paxillae across the arm, at least in smaller specimens, R 40-50 mm, on the two sides of the 
Atlantic. Excluding the two series of superomarginal paxillae, the number in american 
specimens of about this size is proximally c. 13 compared with c. 17 in those from the East 
Atlantic. Unfortunately, owing to their irregular arrangement, it is difficult to make a precise 
estimate of the number of paxillae, especially in larger specimens. Also, the limited amount 
of material available indicates that the paxillar number may increase at a higher rate in 
american specimens. There is a small degree of support for this character in the armament of 
the adambulacral plates. At R c. 40 mm, the third (lateralmost) spine is much smaller than 
the second in american specimens and even at R 70 mm this spine is still much narrower 
basally than the second and only about two-thirds as long. Only when R approaches 
140 mm does the third spine achieve parity in magnitude with the second on most plates. As 
for the accessory adambulacral spinelets, in specimens from both sides of the Atlantic one 
(sometimes two) is usually somewhat enlarged proximal to the third spine (or the space 
between second and third spines). This spinelet is usually one-third to half, but occasionally 
as much as three-quarters as long as the third spine. In american specimens this accessory 
spinelet(s) remains slender but in the eastern Atlantic can be basally stouter and more 
conical in shape. However, there is so much variation in the adambulacral armament that 
these differences cannot be of much importance. Finally, the madreporite may remain 
obscured by the paxillae to a greater extent in specimens from the eastern Atlantic than in 
american ones but this is probably correlated with size (the plate becoming more prominent 
in larger specimens) and is affected by preservation. In total, these minor differences do not 
add up to more than a subspecific distinction between L. elegans and L. sarsi. 

Finally, Koehler (1923) and Carrera-Rodriguez & Tommasi (1977) have recorded 
specimens from Uruguay (33 S) and southern Brazil (c. 30^ S) as L. africana or L. elegans. 
Because of the great geographical discontinuity between them and the rest, comparable to 
that between L. sarsi sarsi and L. sarsi africana, it is not improbable that a minor morpho- 
logical difference justifying a taxonomic distinction of eastern south american specimens 
may exist, most likely again at the subspecific level. 



LECTOTYPE. Because of the complex affinities of Luidia sarsi, it seemed desirable that a 
lectotype be selected from among Sars' material from Bergen to which Diiben & Koren gave 
this name in 1845. Thanks to Dr M. E. Christiansen of the Zoologisk Museum, Oslo, it is 
clear that no type was designated at the time. The specimen chosen (one of seven) was from 
Manger, near Bergen, Oslo Museum reg. no. E1005a, shown in Fig. 6. 

R (maximum)/r is 37/8 mm, = 4-6/1. The shorter arms have R 30 and 32mm. (The 
relatively low R/r ratio is to be expected at this relatively small size.) About 17 lateral 
paxillae correspond to 10 of the more proximal superomarginal paxillae. The median 
paxillae nearly all have a single, slightly coarser, central paxillar spinelet and usually 7-1 1 
peripheral spinelets. Excluding the superomarginal ones, there are c. 17 paxillae across the 
arm breadth basally. The paxillae of the outer lateral row have up to three central spinelets. 

Fig. 6 Luidia sarsi sarsi Diiben & Koren, lectotype, Zoologisk Museum, Oslo reg. no. E1005a, 

Manger, near Bergen. Dorsal view, x 2 - 2. 

180 A.M.CLARK 

All the spinelets of the lateral paxillae are distinctly coarser than those of the median ones, 
while the superomarginal armament is coarser still. The madreporite is not distinguishable. 
The inferomarginal plates bear 2-4 large spines, rather variable in level on the plate, 
sometimes alternating, though at the same level on several consecutive plates. The 
uppermost spine is usually slightly the longest. Most proximal actinal plates bear a single 
blunt, bivalved pedicellaria but no abactinal pedicellariae were observed. Each 
adambulacral bears 3 large spines and an enlarged spinelet one-third to half as long as the 
lateralmost one and proximal to it. Each oral plate bears about four large spines in an 
abradial row and one other spine near the furrow margin, besides a large bivalved 
pedicellaria in the furrow. 

The paralectotypes include four smaller specimens and two larger but broken ones, one 
with R c. 50 mm and the other R/r 55/9*5. The central spinelet of the median paxillae is also 
distinctly coarser than the peripheral ones; c. 20 lateral paxillae correspond to 10 
superomarginal ones; proximally there are usually four inferomarginal spines. 

HOLOTYPE OF Luidia elegans. Because of Perrier's rather inadequate description, this 
specimen was borrowed for reexamination from the Museum of Comparative Zoology, 
Harvard. The number is M.C.Z. 372. It proved to be in fact the specimen illustrated in 
Perrier, 1884, pi. 10, fig. 7, under the name of L. barbadensis, though the negative was 
reversed in printing. One arm is abnormally short, probably regenerated, and curled up at 
the tip. R is c. 40 mm for the longest arm (Perrier gives 35 mm); r is 7 mm; 1 8 lateral paxillae 
correspond to 10 superomarginal ones; proximally there are c. 13 paxillae across the arms 
between (but not counting) the superomarginal ones. The median paxillae mostly have only 
a single central spinelet, not at all coarser than the peripheral ones. No abactinal 
pedicellariae were seen. Most inferomarginal plates bear three large spines, sometimes two, 
showing a tendency for alternation in position. The actinal pedicellariae are bivalved and 
blunt-tipped. The third adambulacral spine is reduced on some plates. The locality is 'Straits 
of Florida, 101 fathoms'. Verrill's guess that Perrier's '101 brasses' meant feet being 

DISTRIBUTION. L. sarsi sarsi from Trondheim Fjord, Norway to Cap Blanc, Mauritania, the 
Azores and the Mediterranean; 9-1 300 metres. 

L. sarsi africana from Luderitz Bay to the Port Elizabeth area of South Africa; 54-360 

L. sarsi elegans from east of New Jersey, U.S.A. (c. 41 N) to the Florida Strait and both 
east and west Gulf of Mexico; also from southern Brazil (c. 30f S) to Uruguay (c. 33 S); 
60-365 metres. There are no reliable records from the Caribbean, Perrier's supposed 
specimens from Barbados being untraced in the M.C.Z. and it is significant that no 
specimens appear to have been taken by the intensive collections of the Atlantis around 
Cuba (H. L. Clark, 1941) or by the Pillsbury in the Caribbean, Lesser Antilles or north of 
South America. 

Luidia senegalensis (Lamarck) 

Asterias senegalensis Lamarck, 1816 : 567. 

Luidia senegalensis: Miiller & Troschel, 1842 : 78, pi. 5, fig. 4; Perrier, 1875 : 342-343 [1876 : 262]; 

Doderlein, 1920 : 249-250, figs 9, 20; H. L. Clark, 1933 : 20-22; Bernasconi, 1943 : 5-6; Tommasi, 

1958:9-11, pi. 2, fig. 2; Ummels, 1963:94-95, pis 10, 11; Downey, 1973:22, pi. 1, figs A, B; 

Blake, 1973 : 30, pi. 2, figs 1-31; Walenkamp, 1976 : 25-29, fig. 5, pi. 1, fig. 4; 1979 : 12, pi. 1, figs 

Luidia marcgravii Steenstrup in Lutken, 1859 : 43^6; Verrill, 1915 : 208-209; Boone, 1933 : 76, pis 

33-36; Bernasconi, 1958 : 125-127. 

REMARKS. A very large specimen from Pillsbury st. 750, off Venezuela, 22-26 metres with R 
up to 252 mm may provide a size record. 
This locally common Caribbean species with its numerous and particularly attenuated 


arms seems even more vulnerable than most Luidias to loss and regeneration of the arm tips, 
probably due to predation, possibly cannibalism. 

DISTRIBUTION. Sporadically in southern Florida and from Jamaica eastwards along the 
Antilles, also Belize and Nicaragua and along the north coast of South America to southern 
Brazil (Santa Caterina, c. 21 S); the original record from Senegal has never been repeated; 
the records from Belize and Nicaragua are new and derived from the Pillsbury collections; 
1-64 metres, the depth range also being extended by a Pillsbury station (745, off Venezuela, 
c. 12 N, 67 W); the previous maximum depth was 45 m off French Guiana. 

Summary of taxonomic changes 

The number of nominal Atlantic species of Luidia is reduced by the present study from a 

one-time maximum of 1 7 to 1 1 . 

Luidia aciculata Mortensen, 19336 is retained as a subspecies of the Indo-West Pacific L. 

sagamina Doderlein, 1920. 

L. africana Sladen, 1 889 is reduced to a subspecies of L. sarsi Diiben & Koren, 1 845. 
L. barimae John & Clark, 1954 is reduced to a subspecies of L. heterozona Fisher, 1940. 
L. bernasconiae A. H. Clark, 1945 is again synonymized with L. alternata (Say, 1825), 

having been revived by Gray, Downey & Cerame- Vivas, 1968. 
L. elegans Perrier, 1 875 is reduced to a subspecies of L. sarsi. 
L. numidica Koehler, 19 1 1 is treated as a subspecies of L. alternata. 
L. quequenensis Bernasconi, 1942 is synonymized with L. alternata. 
L. rosaurae John & Clark, 1954 is synonymized with L. scotti Bell, 1917, which is treated as 

a subspecies of the East Pacific L. ludwigi Fisher, 1906. 


Thanks are due to Miss M. E. Downey of the Smithsonian Institution for access to the Gerda 
and Pillsbury luidiids; also to Dr M. E. Christiansen of the Zoologisk Museum Oslo, and 
Professor R. M. Woollacott of the Museum of Comparative Zoology (M.C.Z.), Harvard, for 
the loan of material. 


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Manuscript accepted for publication 25 June 1 98 1 

New and little known species of Oncaeidae 
(Cyclopoida) from the Northeastern Atlantic 

S. J. Malt 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 


A large number of new species and genera belonging to the Oncaeidae have been described 
in recent years. Many common species have been re-examined and recognized as polytypic, 
comprising two or more varieties (Tanaka, 1960; Moulton, 1973; Ferrari, 1975; Boxshall, 
1977) or two or more species (Heron, 1977). Now that some of the better known species have 
been redescribed in greater detail it is obvious that many of the newly described species are 
also in need of more exact description. Taxonomic study of oncaeids has become a very 
laborious process, demanding the recognition of relatively small morphological differences 
in these microscopic animals, half the known species of which are less than O6 mm in body 

Most records of oncaeids are from the epipelagic zone, but the more recent discovery of 
several species extending down into the deep bathypelagic zone (below 2000 m) indicates 
that previous records reflect the limitations of the sampling programmes rather than any true 
restrictions in their depth range. 

A new species, Oncaea heronae, and two little known species belonging to the genus 
Oncaea are here described. The genera Conaea, Epicalymma and Myctospictosum are 
synonymized with Oncaea. New locality records are presented for six species and two 
previously undescribed males and two juveniles of known species are also described. 

Materials and methods 

The material described below was donated to the British Museum (Natural History) by the 
Institute of Oceanographic Sciences (Wormley) and the Marine Biological Association 
(Plymouth). It was collected in April 1977 at station 9541 (20 N 21 W) in the NE Atlantic 
during cruise 82 of the R.R.S. Discovery, and in September 1979 at the mouth of the English 
Channel (48 N 730' W) during a cruise of the R.V. Sarsia. The Discovery material was 
collected by a 80 jn mesh diatom net attached to the RMT 1 + 8 net system. Sarsia material 
was taken using a pump system with 80 n and 200 // mesh filters. 

Material was preserved in 10% sea water formalin, stained in chlorazol black, dissected in 
lactophenol and mounted in polyvinyl lactophenol. Drawings were made with the aid of a 
camera lucida. Specimens were measured using an eyepiece micrometer, total body length 
being the distance from the tip of the rostrum to the apex of the caudal rami. 

Descriptions of species 
Oncaea tregoubo vi Shmeleva, 1968 
O. tregoubovi Shmeleva, 1968 : 1784-1785, figs 1-12. 

DESCRIPTION Female. Ratio of prosome and urosome lengths 1 -8 : 1 . Prosome elongate, oval 
in dorsal aspect (Fig. la). Mean body length of 3 specimens 0'34 mm (range 0'31 to 

Bull. Br. Mus. not. Hist. (Zool.) 42 (3) : 1 85-205 Issued 27 May 1 982 

186 S. J. MALT 

0*37 mm). Third prosome segment without a dorsal projection. Head (first prosome division) 
1*3 times longer than wide. Proportional lengths of ursome and caudal rami 8 : 57 : 
5:4: 13 : 13 (Fig. Ib). Genital apparatus located on dorsal surface anterior to midpoint of 
genital segment; each area armed with a setule. Maximum width of genital segment at level 
of genital apparatus; length to width ratio 1 '75 : 1 . Caudal rami twice as long as wide. 

First antenna 6-segmented (Fig. Ic); armature: 1-2, II-7, III-3, IV-3, V-2, VI-5 (some 
elements may be missing); ratio of segment lengths 10 : 13 : 46 : 13:8: 10. Second antenna 
3 -segmented (Fig. Id); first segment bearing 1 long pinnate seta distally; second segment 
bearing a row of denticles along internal surface; terminal segment longer than second and 
bearing 5 curved spines and 2 curved setae. Mandible bearing 5 elements (Fig. le): a stout 
seta on external surface, 2 broad blades, 1 bearing a row of setules along internal edge, and 
the other unarmed, and 2 setae, 1 long and hirsute, the other small and unarmed. First 
maxilla bilobed (Fig. 1 f); bearing 1 seta on internal surface; 2 setae on internal lobe and 2 
setae, a setose seta and a spine on external lobe. Second maxilla 2-segmented (Fig. Ig); 
second segment produced distally as an elongate, curved, bilaterally spinulose claw; also 
having an external pinnate seta and an internal bilaterally spinulose element. Maxilliped 
4-segmented (Fig. 1 h); first segment unarmed; internal surface of second segment with 2 
spines, distal spine bidentate; third segment reduced; terminal segment produced as a long 
claw and armed with a row of setules on concave surface, an internal basal spine and an 
external basal setule. 

Endopodites and exopodites of natatory legs 3-segmented (Figs 2a-d); external spines of 
exopodites bilaterally serrate; armature of natatory legs as follows: 

Coxa Basis Endopodite Exopodite 

Legl 0-0 1-1 0-1; 0-1; 0,1, 5 I-0;I-1; III, 1,4 

Leg2 0-0 1-0 0-1; 0-2; II, 1, 3 1-0; !*-!*; Ill, 1, 5 

Leg 3 0-0 1-0 0-1; 0-2; II, 1, 2 1-0; I-1;II,1, 5 

Leg 4 0-0 1-0 0-1; 0-2; II, 1,1 I-0;I-1;II,1, 5 

* = segment missing from figured specimen (Fig. 2b), present in other material. 

Terminal spines of exopodites longer than terminal segments bearing them but terminal 
spines of endopodites shorter than terminal segments; each terminal spine bordered by a 
serrate membrane unilaterally; endopodites of legs 1-3 terminate in conical projections; all 
setae on legs plumose. Fifth leg comprising a small cylindrical segment bearing a single 
terminal seta on body surface near leg (Fig. 1 i). 

MATERIAL EXAMINED. 399 from sample 9541.24, 2980-3560 m. 20 N 21 W. BM(NH) 

REMARKS. This species was identified by the following combination of characters: the 
relative lengths of the segments of the first and second antennae, the armature of the 
maxilliped and natatory legs, and the relative dimensions of the urosome segments. Some 
minor differences were found between the present material and the original material figured 
by Shmeleva (1968). Most notably Shmeleva (1968 : Fig. 9) shows the fourth natatory leg 
with the internal margin of the third endopodite segment unarmed, whereas the present 
specimens have a single seta. The absence of this seta is atypical for Oncaea species. It may 
be significant that Shmeleva does not mention this character in the text of the description, 
and the original drawing may be inaccurate or based on a specimen with incomplete 
armature. This species has only been recorded once before, from the southern Adriatic 
where it was taken between 200 and 300 metres depth. 

Oncaea ivlevi Shmeleva, 1966 

O. ivlevi Shmeleva, 1966 : 932-933, Plate I, figs 1-1 1 . 

DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-7:1 (Fig. 3a). Body length of 

3 specimens 0*33 mm. Third prosome segment without a dorsal projection (Fig. 3b). Head 


1 1 times longer than wide. Proportional lengths of urosome segments and caudal rami 
8 : 50 : 7:6:18:11 (Fig. 3c). Genital apparatus situated anterior to midpoint of dorsal 
surface of genital segment; each area armed with a setule. Maximum width of genital 
segment at the level of genital apparatus; length to width ratio 2:1. Genital segment not 
markedly swollen. Caudal rami nearly twice as long as wide, curving outwards and each 
bearing a stout spine at external corner. 

First antenna armature similar to O. tregoubovi (Fig. 3d); ratio of segment lengths 19 : 19 : 
35 : 12 \ 6 : 9. Second antenna armature similar to O. tregoubovi except that first segment 
armed with a row of setules on internal surface, and second segment with a row of spinules; 
terminal segment equal in length to second segment (Fig. 3e). Mandible and first 
maxilla lost during dissection. Second maxilla like that of 0. tregoubovi (Fig. 30- Maxilliped 
4-segmented (Fig. 3g); first segment unarmed; internal surface of second segment with 2 
spines, longer distal spine spinulose; second segment also bearing a row of denticles along 
internal surface; third segment reduced; terminal claw bearing a row of spinules along 
concave surface and an internal basal spine. 

Armature of natatory legs similar to O. tregoubovi (Figs 3h-k), but terminal segment of 
second endopodite bearing 1 spine on external surface instead of 2; endopodites 2, 3 and 4 
tipped with relatively small conical projections; terminal spines of exopodites 1 and 2 
slightly longer than terminal segments; those of exopodites 3 and 4 slightly shorter; all 
terminal spines of endopodites shorter than terminal segments and bilaterally flanged. 
Fifth leg comprising a small cylindrical free segment bearing 2 setae, outer double length of 
inner; and a seta on body surface near the leg. 

Male. Ratio of prosome and urosome lengths 1-6 : 1 (Fig 4a and b). Mean body length of 6 
specimens - 33 mm (range 0'28 to 0'35 mm). Head I'l times longer than wide. Proportional 
lengths of urosome segments and caudal rami 7 : 60 : 2 : 3 : 2 : 13 : 13. Genital lappets 
produced into small postero-lateral processes (Fig. 4c). Genital segment with length to width 
ratio 2-1 : 1 . Caudal rami twice as long as wide, as in female possessing a stout spine on each 

Mouthparts similar to those of female except first antenna and maxilliped. Three distal 
segments of the first antenna fused. Maxilliped 3 -segmented (Fig. 4d); first segment 
unarmed; second segment bearing 2 short spines and a row of setules on internal surface; 
terminal claw bearing spinules on concave surface and a stout external basal spine. Legs 1-5 
similar to those of female. 

MATERIAL EXAMINED. 399 and 6dtf from sample F166-F170, 8(MO m, 48 N 730' W. 
BM(NH) 99 1981. 115-116,^ 1981. 117-1 19 and 1981. 124. 

REMARKS. This species was identified by the following combination of characters: the 
armature of the natatory legs, the relative dimensions of the urosome segments, and the 
presence of the stout caudal spine. The present specimens differ from Shmeleva's 
descriptions in a few minor details: in the possession of 3 terminal setae in association with 
the spine on the caudal rami rather than 2, the possession of an extra row of spinules on both 
the distal spine of the second segment of the female maxilliped and the terminal claw. There 
are also small differences in the lengths of the terminal exopodite spines. This species has 
also been recorded from the southern Adriatic where it was taken between 50 and 100 metres 
depth (Shmeleva, 1966) and from the Atlantic at 15 stations from 9 S 25 W to 9 S 45 W 
between 10 and 2000 metres depth (Shmeleva, 1969). 

Oncaea hispida (Heron, 1977) Comb. nov. 
Conaea hispida Heron, 1977 : 90-95, Figs 33h-j, 34a-k. 

DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-75 : 1 (Fig. 5a). Body length 
of 1 specimen 0'56 mm. Third prosome segment without dorsal projection. Proportional 
lengths of urosome segments and caudal rami 8:48:7:7: 18: 12 (Fig. 5b). Genital 
apparatus located on dorsal surface anterior to midpoint of genital segment, each area armed 

188 S. J. MALT 

with a setule. Maximum width of genital segment at level of genital apparatus; length to 
width ratio 1'6 : 1. Caudal rami 1*3 times as long as wide. Appendages similar to those 
described by Heron with only minor differences apparent (Figs 5c-h and 6a-d). 

Male. Ratio of prosome and urosome lengths 2 : 1 (Fig. 6e). Body length of 1 specimen 
0'55 mm. Head 1-2 times longer than wide. Proportional lengths of urosome segments and 
caudal rami 10:51 :3:3:3: 18: 12 (Fig.. 6f). Genital lappets produced into small posterior 
processes. Genital segment with length to width ratio 1'5 : 1. Caudal rami approximately as 
long as wide. 

Mouthparts similar to those of female except first antenna and maxilliped. The 3 distal 
segments of first antenna fused. Maxilliped 3-segmented (Fig. 6g); first segment unarmed; 
second segment bearing 2 rows of dentiform processes on internal surface; terminal claw 
unarmed. Legs 1-5 similar to those of female. 

Fifth Copepodid (female). Ratio of prosome and urosome lengths 2 : 1 (Fig 6h). Body 
length of 1 specimen O46 mm. Head approximately as long as wide. Urosome 4-segmented 
(Fig. 6h). Proportional lengths of urosome segments and caudal rami 10 : 45 : 6 : 29 : 10. 
Genital segment without visible genital apparatus, length to width ratio 1'5 : 1. Caudal rami 
1-2 times as long as wide. Appendages similar to those of adult female except that proximal 
seta on second segment of juvenile maxilliped is pinnate (Fig. 6j). 

MATERIAL EXAMINED. 19, Irf, 1 9 juvenile from sample 9541.24, 3980-3960 m, 20 N 21 W. 

REMARKS. This species is here transferred from the genus Conaea to Oncaea (for discussion 
see page 191), and was identified by the following combination of characters: the length of 
the terminal segment of the second antenna and its armature, the armature of the maxilliped 
and the length of the third segment of the fourth endopodite. There are minor differences 
between the present material and Heron's description of the female in the ratio of urosome 
segment lengths, in the proximal spine of the second segment of the maxilliped which lacks 
the spinules in the adult female (but not in the juvenile), and the fifth leg has 2, rather than 1, 
tubercles each bearing at seta. These differences are considered to represent geographical 
variation within the species, as the only previous records of this species are from the antarctic 
zone of the southwest Pacific between 1000 and 2000 metres depth. The male and fifth 
copepodid have not previously been described. 

Oncaea heronae sp. nov. 

DESCRIPTION. Female. Ratio of prosome and urosome lengths 1-5 : 1. Prosome elongate 
and oval in dorsal aspect (Fig. 7a). Mean body length of 4 specimens 0'33 mm (range 0'3 1 to 
0'35 mm). Third prosome segment without a dorsal projection. Head I'l times longer than 
wide. Proportional lengths of urosome segments and caudal rami 9:50:7:7: 13: 14 (Fig. 
7b). Genital apparatus located on dorsal surface anterior to midpoint of genital segment. 
Maximum width of genital segment at level of genital apparatus; length to width ratio 
1 -2 : 1 . Caudal rami twice as long as wide. 

First antenna 6-segmented (Fig. 7c); armature: 1-3, II-6, III-3, IV-1, V-2, VI-5 (some 
elements may be missing); ratio of segment lengths 12 : 22 : 34 : 13 : 7 : 12. Second antenna 
3-segmented (Fig. 7d); first segment bearing 1 long pinnate seta distally; second segment 
unarmed; terminal segment equal to second segment in length, bearing 1 curved spine and 3 
curved setae proximally and 5 curved spines and 1 curved seta distally. Mandible lost during 
dissection. First maxilla (Fig. 7e) and second maxilla (Fig. 70 with armature similar to O. 
tregoubovi. Internal surface of second segment of maxilliped bearing 2 overlapping rows of 
setules and 2 spines; larger distal spine bilaterally dentate (Fig. 7g); terminal claw with 
smooth concave surface; single internal basal spine. 

Armature of natatory legs similar to O. tregoubovi except for armature of terminal 
segments of endopodites 2-4: 


Coxa Basis Endopodite Exopodite 

Legl 0-0 1-1 0-1; 0-1; 0,1, 5 1-0; I-1;III, 1,4 

Leg 2 0-0 ?-0 0-1; 0-2; 1, 1, 3 I-0;I-1;III, 1, 5 

Leg 3 0-0 1-0 0-1; 0-2; 0,1, 2 1-0; 1-1 ; II, I, 5 

Leg 4 0-0 ?-0 0-1; 0-2; 0,1,1 I-0;I-1; II, 1, 5 

Bilaterally serrate spines on external margins of exopodites lack pronounced flanges (Figs 
7h-k); first endopodite with conical terminal projection; all terminal spines shorter than 
terminal segments except terminal spines of fourth exopodite and endopodite, these being 
longer than terminal segments. Fifth leg comprising small free cylindrical segment bearing 2 
terminal setae, the longer, internal seta extending to level of genital apertures. 

Male. Not known. 

MATERIAL EXAMINED. 4 99 from sample 9541.24, 3980-3960 m, 20 N 21W. BM(NH) 
Holotype 1981.125, paratypes 1981.126. 

ETYMOLOGY. This species has been named after Gayle A. Heron of the University of 
Washington, Seattle, in recognition of her work on the taxonomy of the Oncaeidae. 

REMARKS. The second, third and fourth endopodites of O. heronae, O. brodskii Shmeleva, 
1968 and O. longipes Shmeleva, 1968 have the same unusual armature described above, but 
O. heronae can be distinguished by other small differences in the armature of the natatory 
legs, by the 2 overlapping rows of setules on the second segment of the maxilliped, and by the 
structure of the free segment of the fifth leg which is elongate in O. longpipes and reduced to 
a tubercle in O. brodskii, but small and delimited from the body segment in O. heronae. 

Oncaea setosa Heron, 1977 
O. setosa Heron, 1977 : 73, figs 22a-h. 

DESCRIPTION. Female. Mean body length of 4 specimens 0'5 mm (range 0'42 to 0*58 mm). 
Body figured (Figs 8a-c). Appendages of O. setosa similar to those described by Heron with 
only minor differences apparent (Figs 8d-n). 

Male. Ratio of prosome and urosome lengths 1-5:1 (Figs 9a and b). Body length of 1 
specimen 0'58 mm. Head 1-2 times longer than wide. Proportional lengths of urosome 
segments and caudal rami 12 : 47 : 3 : 3 : 3 : 18 : 14 (Fig. 9c). Genital lappets extending 
postero-laterally into acute points. 

Mouthparts like those of female, except first antenna and maxilliped. First antenna 
4-segmented (Fig. 9d). Maxilliped 3 -segmented (Fig. 9e); second segment bearing 2 spines on 
internal surface and 3 groups of setules; terminal claw bearing a stout internal basal spine. 

First and second natatory legs like those of female, but terminal spines of third and fourth 
exopodites longer than those of female, being respectively 1-5 times and twice the length of 
terminal segments. Two spines, representing the fifth leg, borne on tubercle not clearly 
delimited from body. 

Fifth Copepodid (female). Ratio of prosome and urosome lengths 3 : 1 (Figs 9f and g). Body 
length of 1 specimen 0'51 mm. Head 1-2 times longer than wide. Urosome 4-segmented 
(Fig. 9h). Proportional lengths of urosome segments and caudal rami 12 : 40 : 8 : 24 : 16. 
Mouthparts similar to those of adult female except maxilliped, bearing a group of setules on 
internal surface of second segment (Fig. 9i). Natatory legs bearing terminal spines 
proportionally longer than those of adult female (Figs 9j and k, lOa and b). (lOj and k, 1 la 
and b). 

MATERIAL EXAMINED. 499(1 lost), Irf, 1 9 juvenile from sample 954 1.24, 3980-3960 m, 20 N 
21 W.BM(NH) 1981. 127-129. 

REMARKS. This species was identified by the cluster of spinules on the second segment of the 
second antenna, the armature of the maxilliped and the relative dimensions of the urosome 
segments. The female differs from Heron's description in some minor details: the terminal 

190 S. J. MALT 

spines of the third and fourth endopodites are longer in proportion to their terminal 
segments in the present material. The male and the fifth copepodid have not previously been 
described. Heron reported O. setosa from 2 stations in the antarctic zone of the southwest 
Pacific between 1000 and 2000 metres depth. 

Oncaea rotunda Heron, 1977 
O. rotunda Heron, 1977 : 77-79, figs 24o-q, 25a-m. 

DESCRIPTION. Female, body length of 1 specimen - 55 mm. Body (Figs lOc-e) and second 
antenna (Fig. 1 Of) figured. 

MATERIAL EXAMINED. 19 from sample 9541.24. 3980-3960 m, 20 N 21 W. BM(NH) 

REMARKS. This specimen was assigned to O. rotunda because of its distinctive second 
antenna, which has a very short terminal segment relative to the second segment and bears 
relatively short terminal spines and setae. Also the terminal segment of the fourth 
endopodite is reduced. Although somewhat smaller than the type material (mean length 
0'74 mm), this specimen otherwise corresponds closely to Heron's description. Heron 
reported O. rotunda from 3 stations in the antarctic zone of the Pacific between 1000 and 
2000 metres depth. 

Oncaea brocha Heron, 1977 
O. brocha Heron, 1977 : 60, figs 14f-n, 15a-n. 

DESCRIPTION. Female. Body length of 2 specimens 0'81 and 0'83 mm. Urosome (Fig. lOg), 
second antenna (Fig. lOh), maxilliped (Fig. lOi) and second and third natatory legs (Figs lOj 
and k) figured. 

MATERIAL EXAMINED. 299 from sample 9541.24, 3980-3960. 20 N 21W. BM(NH) 

REMARKS. The specimens were identified by the following combination of characters: the 
relative lengths of the segments of the first and second antennae, the armature of the 
maxilliped, of the natatory and of the fifth legs. The second antenna has an additional 
terminal spine not figured by Heron. This may have been lost from Heron's material or may 
represent geographical variation. Another minor difference observed was the length of the 
terminal spines of the second and third endopodites which are relatively longer in the present 
material. The refractive granules mentioned by Heron are absent from these specimens, but 
but these may have been artefacts or may represent the positions of cuticular pores. Heron 
reported O. brocha from 1 station in the antarctic zone of the southwest Pacific between 
1000 and 2000 metres depth. 

Oncaea schmitti (Heron, 1977) Comb. nov. 
Epicalymma schmitti Heron, 1977 : 82-84, figs 28a-n, 29a-e. 

DESCRIPTION. Female. Body length 2 specimens 0*33 and 0'34 mm. Body (Figs 1 la and b), 
second antenna (Fig. 1 1 c) and maxilliped (Fig. 1 1 d) figured. 

MATERIAL EXAMINED. 299 from sample 9541.24, 3980-3960 m. 20 N 21W. BM(NH) 

REMARKS. This species is here transferred from the genus Epicalymma to Oncaea (for 
discussion see page 191). Specimens are clearly identifiable as O. schmitti by the armature 
and relative segment lengths of the second antenna, the armature of the maxilliped and the 
dorsal projections of the caudal rami. However, the specimens are so thinly chitinized that 
the genital segment had partially collapsed during capture or preservation. Therefore, the 
drawing of the urosome may not represent the morphology of the living animal. This species 


has also been recorded from the antarctic zone of the southwest Pacific where it was taken 
between 1000 and 2000 metres depth. 

Oncaea umbonata (Heron, 1977) Comb. nov. 
Epicalymma umbonata Heron, 1977 : 84-86, figs 29f-p, 30a-c. 

DESCRIPTION. Female. Body length of 1 damaged specimen 0*62 mm, without caudal rami. 
Body (Figs 1 1 e and 0, second antenna (Fig. 1 1 g) maxilliped (Fig. 1 1 h) figured. 

MATERIAL EXAMINED. \g from sample 9541.24, 3980-3960 m. 20 N 21 W. BM(NH) 

REMARKS. This species is here transferred from the genus Epicalymma to Oncaea (for 
discussion see below). Although the specimen lacks its caudal rami it was assigned to O. 
umbonata on the basis of its body size and maxilliped armature. There are also differences of 
the armature of the second antenna between this and the specimens recorded as O. schmitti, 
the two species are otherwise very close. The natatory legs and other mouthparts correspond 
closely to Heron's original description. This species has also been recorded from the 
antarctic zone of the southwest Pacific where it was taken between 1000 and 2000 metres 

Phylogenetic relationships within the genus Oncaea 

Giesbrecht (1892) designated the following character states as typical of the genus 
Conaea: the elongate terminal segment of the second antenna with very large, hook-tipped 
spines; endopodite of the fourth leg shorter than the exopodite, third segment is reduced 
(shorter than the first or second segment); the fifth leg reduced to a single plumose seta. 
However Heron (1977) was able to demonstrate the presence of a minute spinule 
accompanying the seta of the fifth leg of O. gracilis Dana, 1852 (=syn. C. rapax), the type 
species. Heron also described 2 new species of Conaea, C. succurva and C. hispida. Both of 
these have a seta on a small prominence representing leg 5, and a reduced endopodite of the 
fourth leg, but the third endopodite segment exceeds the first segment in length. These 2 
species also lack the hooked tips to the terminal spines of the second antenna. Oncaea 
expressa Gordejeva, 1973 shares these and other character states but differs in the 
morphology of the natatory legs (Table 1). No less than 27 other species of Oncaea also show 
1 or more of these Conaea-\ike character states (Table 2) The 2 species of Epicalymma 
which Heron (1977) described also exhibit the following character states: elongate terminal 
segment of the second antenna with very large hook-tipped spines, endopodite of the fourth 
leg shorter than the exopodite, third segment reduced (but longer than the first or second 
segment), fifth leg with a single terminal seta. 

A continuous gradation of character states within Oncaea, Conaea and Epicalymma is 
thus becoming apparent as more species of Oncaea are discovered. The armature of the 
third exopodite segments of the swimming legs also illustrates this. Most Oncaea species 
have an external spine formula of III, III, II, II but there is-considerable variation within the 
genus: II, III, III, I (O. ancora Gordejeva, 1973); III, III, III, II (O. minor Shmeleva, 1979 and 
O. parobscura Shmeleva, 1979); III, II, II, I (O. expressa Gordejeva. 1973); III, II, II, II (O. 
brodskii Shmeleva, 1968 and O. longipes Shmeleva, 1968); II, III, II, I (O. exigua Farran, 
1908); II, II, I, I (O. atlantica Shmeleva, 1967 and O. vodjanitskii Shmeleva and Delalo, 
1969; II, III, II, II (O. zernovi Shmeleva, 1966 and O. mollicula Gordejeva, 1975). The 
typical formula for Conaea is II, III, II, I and for Epicalymma II, III, III, I. Such variation 
represents a continuum which cannot, in my opinion, justifiably be subdivided into 2 or 
more genera as it is at present. It is probable that Conaea-\\ke and Epicalymma-like 
character states have been derived by reduction and loss of armature elements from ancestors 
within the genus Oncaea. Separation of the Conaea and Epicalymma species would leave 

192 S. J. MALT 

Table 1 Comparison of morphology of natatory legs ofOncaea expressa. O. succurva and O. hispida. 


Leg 2 

Leg 3 

Leg 4 

O. expressa 

Exopod. spine 

Exopod. spine 

Exopod. spine 

Exopod. spine 

Gordejeva, 1973 

2 x length 

2 x length 

2- 5 x length 

> 2x length 

term. segm. 

term. segm. 

term. segm. 

term. segm. 

Endopod. spine 

Endopod. spine 

Endopod. spine 

Endopod. spine 

shorter than 

1-5 x length 

>4x length 

> 4 x length 

term. segm. 

term. segm. 

term, segm 

term. segm. 

O. succurva 

Exopod. spine 

Exopod. spine 

Exopod. spine 

Exopod. spine 

(Heron, 1977) 

2 x length 

>2 x length 

2-5 x length 

>2'5 x length 

comb. nov. 

term. segm. 

term. segm. 

term. segm. 

term. segm. 

(syn. Conaea 

Endopod. spine 

Endopod. spine 

Endopod. spine 

Endopod. spine 


same length 

1-5 x length 

2- 5 x length 

3 x length 

term. segm. 

term. segm. 

term. segm. 

term. segm. 

O. hispida 

Exopod. spine 

Exopod. spine 

Exopod. spine 

Exopod. spine 

(Heron, 1977) 

2 x length 

>2x length 

2-5 x length 

> 3 x length 

comb. nov. 

term. segm. 

term. segm. 

term. segm. 

term. segm. 

(syn Conaea 

Endopod. spine 

Endopod. spine 

Endopod. spine 

Endopod. spine 


same length 

1 -5 x length 

2 x length 

>4 x length 

term. segm. 

term. segm. 

term. segm. 

term. segm. 

Oncaea as a paraphyletic group. In order to retain Oncaea as a monophyletic group (sensu 
Hennig, 1966) it is proposed that Conaea and Epicalymma be synonymized with Oncaea 
and that the species of Conaea and Epicalymma be transferred to Oncaea, C. gracilis, C. 
succurva, and C. hispida becoming O. gracilis, O. succurva and O. hispida, also E. schmitti 
and E. umbonata becoming O. schmitti and O. umbonata respectively. 

Kazatchenko and Andreev (1977) described Myctospictosum as a genus of uncertain 
taxonomic position. A single male of M. philippinensis was found on the gills ofMyctophum 
spinosum in the western Pacific. It is clear from the figures (Kazatchenko and Andreev, 
1977 : Figs 9 and 10) that this is an oncaeid. It has the body shape, the 4-segmented first 
antenna and the 3-segmented second antenna of a typical Oncaea male. The other 
mouthparts have been interpreted differently by the authors but are clearly those of a male 
Oncaea. The natatory legs are also typical of the genus. It is proposed, therefore, that 
Myctospictosum should be synonymized with Oncaea. The armature of the natatory legs and 
the relative lengths of the posterior segments separate O. philippinensis comb. nov. from any 
of the known males of the genus Oncaea. 

Oncaeids are often found clinging by their maxillipeds to other organisms and debris in 
plankton samples, there is, therefore, no reason to suppose that O. philippinensis is parasitic, 
it is probably a free-living planktivore like other species of the genus. 

The generic diagnosis ofOncaea is ammended as follows: 

ONCAEA Philippi, 1843 

Oncaea Philippi, 1843 : 63. 

AntariaDana, 1846:229. 

Conaea Giesbrecht, 1891 : 477. 

Epicalymma Heron, 1977 : 82. 

Myctospictosum Kazatchenko and Andreev, 1977 : 47. 

DIAGNOSIS. Cyclopoid shape. Rostral area thickened with rounded posteroventral margin. 
Female urosome 5-segmented, male 6-segmented. First antenna 6-segmented in female, 


Table 2 Conaea-like character states exhibited by species ofOncaea. 

c rj 

u . o 

r; CO O 

SP C . u Q C M-5 p 

?s ^ U^ -*-^ <U I < ,._ 

6CO tJ -4 > C QjTO CO ^^ 

-C J= - C *> c P.^c 

So c o o 

- o o 

B - * 2 

*><=; ^ -o 3 c ^TO> 

l|s || -g|^ 1| l"8-i 

PCoo mj^t- SCtu o w e 

8 I ^^ 'o 8 g B| a 

Ha><u W*j Jxi5 m2^'Z^ H ' o G c/5 

O. graci/w (Dana, 1852) 

O. schmitti (Heron, 1977) 

O. umbonata (Heron, 1977) 

O. ancora Gordejeva, 1973 

O. atlantica Shmeleva, 1967 

O. exigua Farran, 1908 

O. expressa Gordjeva, 1973 

O. hispida (Heron, 1977) 

O. mollicula Gordejeva, 1975 

O. succurva (Heron, 1977) 

O. alboranica Shmeleva, 1979 

O. prendeli Shmeleva, 1966 

O. shmelevi Gordejeva, 1972 

O.tenellaSars, 1916 

O. tregoubovi, Shmeleva, 1968 

O. brodskiiShme\e\&, 1968 

O. heronae sp. nov. 

O. longpipes Shmeleva, 1968 

O. memorata Gordejeva, 1973 

O. minima Shmeleva, 1968 

O. rotunda Heron, 1977 

O. rotundata Boxshall, 1977 

O. subtilis Giesbrecht, 1 892 

O. vodjanitskii Shmeleva & Delalo, 1969 

O. zernovi Shmeleva, 1966 

O. ornata Giesbrecht, 1 89 1 

O. infantula Gordejeva, 1972 

O. africana Shmeleva, 1979 

O. curvata Giesbrecht, 1902 

O. englishi Heron, 1977 

O. ivlevi Shmeleva, 1966 

4-segmented in male, both with long third segment. Second antenna 3-segmented, terminal 
segment bearing proximal group of 3 or 4 setae and a distal group of 6 to 8 setae. Mandible 
bearing 2 blades and 2 or 3 setae. First maxilla bilobed, inner lobe with 3 elements. Second 
maxilla with setose claw. Maxilliped 4-segmented in female and 3-segmented in male, 
second segment in female bearing 2 setae on the internal surface. Terminal segment 
produced as a long, curved claw. Legs 1-4 3-segmented, endopodites slender. Exopodites 
armed with serrate spines *on external margin. Leg 5 reduced to small rod or knob-like 
segment fused to urosome somite and bearing 1 or 2 apical setae. Gender feminine. 

TYPE SPECIES: Oncaea vensusta Philippi, 1843. Sixty-eight species of Oncaea are now 
known, over half of which have been described within the past 15 years (Shmeleva, 1966, 
1967, 1968, 1969, 1979; Razouls, 1969; Gordejeva, 1972, 1973, 1975; Boxshall, 1977; 
Heron, 1977; Kazatchenko and Andreev, 1977). 

194 S. J. MALT 


I wish to thank Dr H. S. J. Roe (I.O.S.) and Dr R. P. Harris (M.B.A.) for arranging for 
material to be donated to the BM(NH), also Dr G. A. Boxshall and Dr R. J. Lincoln for 
reading and commenting on the manuscript. This work was undertaken while the author was 
in receipt of a BM(NH) postgraduate studentship. 


Boxshall, G. A. 1977. The planktonic copepods of the northeastern Atlantic Ocean: some taxonomic 
observations on the Oncaeidae (Cyclopoida). Bull. Br. Mus. nat. Hist. (Zool.) 31 (3) : 101-1 55. 

Dana, J. D. 1 846. Notice of some genera of Cyclopacea. Am. J. Sci. ser. 2, 1 : 225-230. 

1852. Crustacea. U.S. Explor. Exped. 13 : 1-1618. 

Ferrari, F. D. 1975. Taxonomic notes of the genus Oncaea (Copepoda : Cyclopoida) from the Gulf of 
Mexico and northern Caribbean Sea. Proc. biol. Soc. Wash. 88 (2 1 ): 2 1 7-232. 

Giesbrecht, W. 1891. Elenco dei Copepodi pelagici raccolti dal tenente di vascello Gaetano Chierchia 
durante il viaggio della R. Corvetta 'Vettor Pisani' negli anni 1882-1885, e dal tenente di vascello 
Francesco Orsini nel Mar Rosso, nel 1 884. Alt. Acad. naz. Lincei Re. 7 (10) : 474^8 1 . 

1892. Systemik und Faunistik des pelagischen Copepoden des Golfes von Neapel und der 

angrenzenden Meeres-Abschnitte. Fauna Flora Golf. Neapel 19 : 1-83 1 . 
1902. Copepoden. Result. Voyage S. Y. Belgica, Zool. 1-49. 

Gordejeva, K. T. 1972. New species of Oncaea (Copepoda, Cyclopoida) from the tropical zone of the 

Atlantic Ocean. Zool. Zh. 51 (7) : 963-968. (In Russian). 
1973. New species of the genus Oncaea (Cyclopoida) from the tropical Atlantic. Zool. Zh. 

52(10): 1572-1 576. (In Russian). 

1975. Pelagic Cyclopoida (Copepoda) from the tropic Atlantic and South Seas. Zool. Zh. 

54 (5): 776-779. (In Russian). 
Hennig, W. 1966. Phylogenetic systematics. Univ. of Illinois Press, Urbana. 263 pp. 
Heron, G. A. 1977. Twenty-six species of Oncaeidae (Copepoda : Cyclopoida) from the southwest 

Pacific-Antarctic area. Biology of the Antarctic Seas. VI. Antarctic Res. Ser. Washington 26 : 37-96. 
Kazatchenko, V. N. & Andreev, G. V. 1977. Parasitic copepods (Crustacea) collected during 57th 

cruise of 'Vityaz' in the western tropical Pacific and seas of the Indo-Malayan archipelago. Trudy 

Inst. Okeanol. 107 : 30^8. (In Russian). 
Moulton, T. 1973. Principal component analysis of variation in form within Oncaea conifera 

Giesbrecht 1 89 1 , a species of copepod (Crustacea). Syst. Zool. 22(2) : 141-156. 
Philippi, A. 1843. Fernere Beobachtungen iiber die Copepoden des Mittelmeeres. Arch. Naturgesch. 

9(1): 54-71. 
Razouls, C. 1969. Description d'une espece nouvelle du genre Oncaea (Copepoda,, Cyclopoida). Vie 

Sars, G. O. 1916. Liste systematique des Cyclopoides, Harpacticoides et Monstrilloides recueillis 

pendant les campagnes des S.A.S. le Prince Albert de Monaco, avec descriptions et figures des 

especes nouvelles. Bull. Inst. oceanogr. Monaco. 323 : 1-15. 
Shmeleva, A. A. 1966. New species of the genus Oncaea (Copepoda, Cyclopoida) from the Adriatic 

Sea. Zool. Zh. 45 (6) : 932-936. (In Russian). 

1967. New Oncaea species (Copepoda, Cyclopoida) from south-western part of the Atlantic 

Ocean. Zool. Zh. 46 (4) : 62 1-622. (In Russian). 

1968. New species of planktonic Copepoda : Cyclopoida from the Adriatic Sea. Zool. Zh. 

47(12): 1784-1 793. (In Russian). 

1969. Especes nouvelles du genre Oncaea (Copepoda. Cyclopoida) de la mer Adriatique. Bull. 

Inst. oceanogr. Monaco 68, No. 1393 : 1-28. 

1979. New species and some previously unknown males of the genus Oncaea from the 

Mediterranean. Zool. Zh. 58 (4) : 491^98. (In Russian). 
Tanaka, O. 1960. Pelagic Copepoda. Spec. Publs. Seto mar. Biol. Lab. 10 : 1-177. 

Manuscript accepted for publication 4 June 1 98 1 



Fig. 1 Oncaea tregoubovi. a. female, dorsal; b. urosome, dorsal; c. first antenna; d. second 
antenna; e. mandible; f. first maxilla; g. second maxilla; h. maxilliped; i. fifth leg. 





Fig. 2 Oncaea tregoubovi. a. female, leg 1 ; b. leg 2 (damaged); c. leg 3; d. leg 4. 



Fig. 3 Oncaea ivlevi. a. female, dorsal; b. female, lateral; c. urosome, dorso-lateral; d. first 
antenna; e. second antenna; f. second maxilla; g. maxilliped; h. leg l;i. Ieg2;j. leg 3; k. leg 4. 







Fig. 4 Oncaea ivlevi. a. male, dorsal; b. male, lateral; c. urosome, lateral; d. maxilliped. 



Fig. 5 Oncaea hispida. a. female, lateral; b. urosome, dorsal; c. first antenna; d. second antenna; 
e. mandible; f. second maxilla; g. maxilliped; h. labrum (lamella missing). 



Fig. 6 Oncaea hispida. a. female, leg 1 ; b. leg 2; c. leg 3; d. leg 4; e. male dorsal; f. urosome ventral 
(specimen distorted); g. maxilliped; h. female copepodid V, dorsal; i. posterior segments of 
urosome, dorsal (only posterior half of genital segment figured);], maxilliped. 




Fig. 7. Oncaea heronae sp. nov. a. female, dorsal; b. urosome, dorsal; c. first antenna; d. second 
antenna; e. first maxilla, f. second maxilla; g. maxilliped; h. leg l;i. leg 2; j. leg 3; k. leg 4. 



Fig. 8 Oncaea setosa. a. female, dorsal; b. female, lateral; c. urosome, dorsal; d. first antenna; e. 
second antenna; f. mandible; g. first maxilla; h. second maxilla; i. maxilliped; j. labrum (lamella 
missing); k. leg 1 ; 1. leg 2; m. leg 3; n. leg 4. 



Fig. 9 Oncaea setosa. a. male, dorsal; b. male lateral; c. urosome, ventral; d. first antenna; e. 
maxilliped (distal spine missing); f. female copepodid V, dorsal; g. female copepodid V, lateral; 
h. urosome, dorso-lateral; i. maxilliped; j. leg l;k. leg 2. 



Fig 10 Oncaea setosa. a. female copepodid V, leg 3; b. leg 4; Oncaea rotunda, c. female, dorsal; d. 
female, lateral; e. urosome, dorsal; f. second antenna; Oncaea brocha. g. female urosome, dorsal; 
h. second antenna; i. maxilliped;j. leg 2; k. leg 3. 



0-1 mm 



Fig 11 Oncaea schmitti. a. female, dorsal; b. female, lateral; c. second antenna; d. maxilliped; 
Oncaea umbonata. e. female, dorsal; f. female, lateral; g. second antenna; h. maxilliped. 

Larval and post-larval development of the 
Slender-legged Spider Crab, Macropodia rostrata 
(Linnaeus) (Oxyrhyncha : Majidae : Inachinae), 
reared in the laboratory 

R. W. Ingle 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 


The Slender-legged Spider Crab Macropodia rostrata (Linnaeus) occurs northward to about 
latitude 65 N and southward to the Moroccan coast and Mediterranean (see Christiansen, 
1969; Monod, 1932; Zariquiey Alvarez, 1968). West African and Cape Verde Island 
specimens have been assigned recently to M. spinulosa (Miers) (see Manning & Holthius, 
1981). M. rostrata has been reported also from False Bay, S. Africa (see Barnard, 1950). 

Previous descriptions (see p. 207) of larval stages of M rostrata are not sufficiently detailed 
for comparative studies. In 1974 larvae of this species were reared from crabs collected from 
Tunisian and Plymouth waters and in 1977 from specimens obtained off the Isle of Man. 
These rearings provided material for the present detailed description and comparisons of the 
larvae and first crab stages of M. rostrata from three localities within its range of distribution. 

Materials and methods 

Ovigerous crabs were trawled from the following localities. (1) Off Port Erin, Isle of Man. 
20-24 m., 24-25.3.1977. (2) S.W. of Eddystone Lighthouse, nr. Plymouth, Devon, 90 m., 
1 8.6. 1 974. (3) OffCarthage-Salammbo, Tunisia, 4-5 m., 20.2. 1 974. 

Larvae were reared using methods described by Rice & Ingle (1975 : 104) and Ingle & 
Clark (1977). Material was fixed in Steedman's fixative (Steedman, 1976 : 148) and later 
transferred to 70% alcohol. Drawings and measurements were made with the aid of a camera 
lucida. Measurements given are: (a) distance from base to tip of dorsal spine (D.S.); (b) 
carapace length, from between eyes to posterio-lateral carapace margin (C.L.). 

All material has been incorporated into the Collections of the British Museum (Natural 
History), accession no;- 1981:201-225. Drawings and descriptions (unless otherwise 
indicated) are of specimens from locality (1). Ten specimens of each stage were dissected and 
examined from each locality except for ZII of Tunisian material of which 20 specimens were 
dissected to confirm the absence of a seta on the outer margin of the maxillule basis (see 
p. 2 10). 

Whilst in the rearing laboratory females and larvae were maintained at 15C. Larvae from 
locality (1) hatched within 10-12 days and those from (2) within 6-7 days of collecting the 
females. Both hatchings took an average of 30 days to reach first crab stage. Those from 
locality (3) hatched within 14 and 63 days respectively from time of collecting and an 
average of 22 days elapsed before the appearance of first crab stage. 



Larval references. Macropodia phalangium:- Thompson, 1 836 : 37 1 , fig. e (pre-1 st zoea), non fig. 2 = ? 

Bull. Br. Mus. nal. Hist. (Zool.) 42 (3) : 207-225 Issued 27 May 1982 

208 R. W. INGLE 

Hyas sp. ? Stenorhynchus rostrata:- Stuxberg. 1 874 : 3 ( 1 st zoea). ? Stenorhynchus phalangium:- Cano, 
1893:Tav. 35, figs 70, 72, 74, 77-85 (1st zoea, megal. crab). Macropodia rostrata:- Labour, 
1928 : 550, PI. Ill, fig. 10 (coloured), PI. XV, fig 7, PI. XVI, figs 3, 5, 8 (prezoea, 1st, 2nd zoeae, megal. 
1st crab), non Stenorhynchus phalangium:- Lo Bianco, 1904 : 439; 1904 : Taf. 12, fig 43 (=? Inachus 


Dimensions: D.S. 1-3-1-4 mm.,C.L.O-7-0-8 mm. 

Carapace (Fig. la). Dorsal spine long and straight, narrowing distally. Rostral and lateral 

spines not developed. Dorso-median elevation conspicuous above which there is a small seta 

on either side; ocular eaves developed; a pair of small dorso-lateral setae near base of dorsal 

spine; posterio-lateral margin of carapace with 'majid' spine and 1-2 additional long thin 

setae (see inset Fig. la). Surface of carapace with minute spinules. 

Eyes: Partly fused to carapace. 

Antennule: (Fig. 2a). Unsegmented and with 2 terminal aesthetascs and 2 setae. 

Antenna: (Fig. 2a). Spinous process without distal spinules: exopod slightly shorter than 

spinous process and with one small spinule in proximal half; endopod developed as acute 


Mandible: (Fig. 2b). Incisor and molar process developed, palp absent. 

Maxillule: (Fig. 3a). Endopod 2-segmented (incipiently in some specimens), distal segment 

long and thin with 3 long distal setae; basal endite with 5 spines and 2 setae, coxal endite with 

7 setae/spines. 

Maxilla: (Fig. 4a). Endopod narrow, distally truncate and with 4 long setae; basal endite 

broad, distal margin concave or incipiently bilobed and with 4 + 5 setae; coxal endite 

incipiently bilobed, outer lobe with acute outer margin, with 3+4 setae respectively; 

scaphognathite with 10 long plumose marginal setae, posterior seta very stout. 

First maxilliped: (Fig. 5a). Basis with 9 setae arranged 2, 2, 2, 3; endopod 5 -segmented with 

3,2,1,2,4 + 1 setae; exopod with 4 terminal natatory setae. 

Second maxilliped: (Fig. 5b). Basis with one proximal seta; endopod 3-segmented with 0, 0, 

3-4 + 1 setae; exopod incipiently (in some specimens) segmented with 4 terminal natatory 


Third maxilliped and pereiopods: Represented as incipient buds. 

Abdomen: (Figs 6a, b, e). 5-segmented + telson; 2nd segment with a pair of anterio-laterally 

directed acute dorso-lateral processes; posterio-lateral processes on segments 3-5 long and 

terminally acute; a pair of minute setae near posterio-dorsal margins of segments 2-5. Telson 

furcae long and distally with minute spinules, each furca with a small lateral spine; inner 

medio-lateral margin of telson with 3 long plumed setae, innermost (in some specimens) 

noticeably short (see Fig. 6e); incipient pleopod buds on segments 2-5. 


Dimensions: D.S. 1-1-1-2 mm., C.L. 0-8-0-9 mm. 

Carapace (Figs Ib, c). Dorsal spine curved. Dorso-median elevation well developed; ocular 

eaves conspicuously expanded (Fig. Ic). 

Eyes: Moveable. 

Antennule: (Fig. 2c). With 6 aesthetascs and 1-2 setae. 

Antenna: (Fig. 2c). Exopod with 2 proximal spinules; endopod slightly more than \ length of 


Mandible: (Fig. 2d). Each half dissimilar. 

Maxillule: (Fig. 3b). Outer margin of basal endite with a seta, distal margin with 5 spines and 

3 setae; coxal endite with 7 setae/spines. 

Maxilla: (Fig. 4b). Endopod setation unchanged; basal endite with 5 + 5 setae; coxal setation 

unchanged; scaphognathite with 16 setae. 

First maxilliped: (Fig. 5c). Setation on basis and endopod unchanged; exopod with 6 

terminal natatory setae. 


Second maxilliped: (Fig. 5d). Basis without setae; endopod with 0, 0, 3 + 1 setae; exopod 

with 6 terminal natatory setae. 

Third maxilliped and pereiopods: Represented as conspicuous incipient buds. 

Abdomen: (Fig. 6c, d). Posterio-lateral processes on segments 3-5 longer than in first stage; 

pleopod buds long. 


Dimensions: C.L. 1 '2-1*3 mm. 

Carapace: (Figs Id, e). Longer than broad, with a small obtuse rostrum and obtuse but 

prominent submedian lobes; frontal region with 2-3 median broad longitudinal tubercles, 

hepatic regions swollen, each protogastric region with a long anteriorly directed process 

narrowing distally; cardiac region with a posterio-dorsally directed spine. 

Eyes: Large, with well developed cornea. 

Antennule: (Fig. 2e). Peduncle 3-segmented, terminal segment with a seta on inner margin; 

exopod 2-segmented, proximal with 2 and distal segment with 4 aesthetascs. 

Antenna: (Fig. 2f). Peduncle 3 segmented, proximal segment with a stout ventrally directed 

process, distal segment with a small ventral seta; flagellum 4-segmented, 2nd segment longest 

and with 4 setae, 4th segment with 2-3 aesthetascs. 

Mandible: (Fig. 2g). Molar process slightly reduced, incisor prominent and broad; palp stout 

and unsegmented, with a small terminal seta. 

Maxillule: (Fig. 3c). Endopod reduced; distal margin of basal endite with 6-7 spines and 6-7 

setae, coxa with 6-7 setae/spines. 

Maxilla: (Fig. 4c). Endopod reduced and terminally acute, with one long distal seta; basal 

endite broadly oval, with 6 + 3-4 setae, coxal endite with 3 + 3 setae; scaphognathite with 

18-1 9 setae. 

First maxilliped: (Fig. 7a). Coxa with 4, basis with 9-10 setae; exopod 2-segmented, distal 

segment with 4 setae; endopod reduced and terminally acute; epipod small. 

Second maxilliped: (Fig. 7b). Exopod 2-segmented, proximal segment long, distal with 4 

setae; endopod 4-segmented, 2nd (carpus) with one, 3rd (propodus) with 3 setae and 4th 

(dactylus) with one seta and 3 spines; epipod small (not shown in figure). 

Third maxilliped: (Fig. 8a). Endopod 5-segmented, inner margin of ischium with few minute 

spinules and 7-9 short setae, merus with 4 setae, carpus with 3, propodus with 6 and dactylus 

with 4 setae; distal segment of exopod with 4 setae, epipod moderately well developed. 

Pereiopods: (Figs 8c, 9a, lOa-c). Chelipeds moderately stout and sparsely setose, distal parts 

of propodus and dactylus curved; merus with small obtuse spinule on inner proximal 

margin; basis/coxa with small obtuse process on lower margin. Pereiopods 2-5 slender, 

setose and minutely spinulate, dactylus terminally acute, ischium of 2nd-3rd pereiopods 

with a prominent curved hook-shaped process. 

Abdomen: (Figs If, g). 5-segmented + telson; 1st with 4, 2nd-3rd with 2 posterio-dorsal setae, 

4th with 4 setae, 5th with 6 setae; telson broader than long, posterior margin subtruncate. A 

pair of well developed pleopods on segments 2-5, 4th pair smallest, exopod of each with 8 

long plumose setae; endopod of each with 2 coupling hooks. 


Dimensions: C.L. 1 -4-1-5 mm. 

Carapace: (Fig. Ij). Slightly less that lx longer than broad; frontal region slightly 

produced, submedian lobes obtuse and with clusters of hook-setae, each protogastric region 

with a tubercle; orbits long, hepatic regions swollen, epibranchial regions slightly expanded, 

mesogastric with 2 median cristate tubercles, cardiac region with a prominant tubercle, 

intestinal with a small median tubercle; dorsal surface and lateral margins of carapace with 

many hook-setae. 

Eyes: Large and with a few hook-setae. 

Antennule: (Fig. 2h). First and 3rd segments of peduncle with a seta; exopod indistinctly 

segmented, with 7-8 aesthetascs and 2-3 setae; endopod 2-segmented, with 2 terminal and 2 

sub-terminal setae. 

210 R. W. INGLE 

Antenna: (Fig. 2h). Peduncular segments with 2, 1-2, and 5 setae respectively; flagellum 

usually with 3 or 4 demarcated segments, with 3 subterminal setae as shown. 

Mandible: (Fig. 2i). Molar process acute, incisor expanded as a broad cristate lobe; 

mandibular palp 2-segmented, distal segment with 2 setae. 

Maxillule: (Fig. 3d). Endopod very reduced, outer margin of basal endite with a prominent 

seta, margin with 8 spines and 4 setae; coxal endite with 9 setae/spines. 

Maxilla: (Fig. 4d). Endopod very reduced and with one apical seta; basal endite with 10 and 

coxal with 2-3 setae respectively; scaphognathite with 20-2 1 short plumose setae. 

First maxilliped: (Fig. 7c). Coxa with 10-11 setae, basis with 10 setae; distal segment of 

exopod with 3 long setae; endopod sub-triangular, with 1-2 small setae on distal margin; 

epipod well developed and with 6 setae. 

Second maxilliped: (Fig. 7d). Merus with 2 setae, propodus with 4 setae and a distal spine, 

dactylus with 1-2 setae and 4 spines; epipod small. 

Third maxilliped: (Fig. 8b). Ischium with numerous setae (18-20) as shown, inner margin 

with 3-4 processes; merus with 3 setae, inner margin with an acute process; carpus, propodus 

and dactylus with 2, 4 and 5 setae respectively on outer surface and margins; distal segment 

of exopod with 3 short plumose setae; a cluster of short setae near coxal/epipod junction; 

epipod well developed. 

Pereiopods: (Figs 8d, 9b, c, lOd, e). Cheliped moderately setose as shown; inner distal 

propodal margin with 3 acute to sub-acute teeth (see inset to Fig. 8d); distal part of the 

propodus acute and curved. Pereiopods 2-5 long, thin and with numerous setae; 2nd and 3rd 

similar in shape, 3rd longest (Fig. 9c); 4th and 5th similar in shape, dactylus curved, inner 

margin of dactylus of 4th (Fig. lOd) with 4 spines, of 5th with 2-3 spines. 


Comparisons of larvae and first crab stages of Mediterranean and British material of M. 
rostrata (see p. 207) have revealed only two apparent morphological differences. (1) all zoeae 
II examined from the Tunisian locality are without a seta on the outer margin of the 
maxillule basal endite; this seta is present in all zoeae II reared from the Plymouth and Isle of 
Man crabs. The presence of this seta is considered an important larval brachyuran feature 
(see Rice, 1980 : 299 as 'exopod' seta), and its absence in the Tunisian specimens cannot be 
explained. Clark (1980) also reported its absence from the maxillule of zoeae II Inachus 
dorsettensis (Pennant) reared from Isle of Man crabs but found it present in Plymouth 
material. (2) in the first crab stage of Tunisian specimens the carapace submedian lobes are 
widely spaced and the outline of the hepatic region is noticeably convex. By comparison, 
first crab stages from the two British localities are slightly smaller than the Tunisian speci- 
mens, the submedian lobes are closer together and the hepatic region outline is relatively 
straight (Fig. 1, cf j & k). These slight morphological differences observed in the first crab 
stages may express the extremes of geographical variation of M. rostrata since the 
Mediterranean material probably represents the near southernmost limit of this species (see 
p. 207). Ingle & Manning (in press) have drawn attention to the noticeable variation of 
carapace shape among population of pre- and post pubertal crabs of M. rostrata from the 
N.E. Atlantic and Mediterranean regions. 

Four species of Macropodia are reported from N.E. Atlantic waters: M. rostrata 
(Linnaeus), M. deflexa Forest, M. tenuirostris (Leach) and M. linaresi Forest & Zariquiey 
Alvarez. Larval stages of the first three species mentioned were described briefly by Lebour 
(1927, 1928); larvae ofM. linaresi are unknown. 

Lebour (1927, 1928) recognized that the zoeae of M. rostrata and M. deflexa (=egyptia) 
has a longer and more straight dorsal spine, and longer antennae than M. tenuirostris 
(= longirostris); she also noted that the antennae of M. deflexa are longer than M. rostrata. 
Lebour remarked that the megalope of M. rostrata has shorter legs and a less deflected 
rostrum than M. tenuirostris, and that the 'central lobe' of the rostrum of M. deflexa 


protrudes more than that of M. rostrata but that otherwise the megalop of both species are 
'hardly to be distinguished'. 

Larval material of M. dejlexa has not been available for examination during this study but 
it has been possible to compare the larvae of M. rostrata with those of M tenuirostris reared 
from females collected off the Isle of Man during March 1977. These comparisons are listed 
in Table 1 . 

Table 1 


M. rostrata 

M. tenuirostris 

Zoea I 

Carapace length: 
Dorsal spine length: 

Antenna, total length: 
Second mxpd endop. 3rd sgmt: 

Zoea II 

Carapace length: 
Antenna, total length: 

Carapace dorso-median 

Maxilla, coxal endite: 

Abdomen posterio-lateral 

Second mxpd endop. 3rd sgmt: 

Carapace length: 
Carapace frontal region: 

Carapace protogastric 

Cheliped merus: 

0-7-O8 mm 
1*3-1-4 mm 
relatively straight 
1-2-1 -2 mm 
4 + 1 setae 

0-8-0-9 mm 
1'3-1'4 mm 
(exceeding C.L.) 

average of 7 setae 
average of 1 6 setae 


3-4 + 1 setae 

average 1 -2 mm 
2-3 median longit. 

forming moderately wide 

with a small spine but 
without secondary pos- 
terior spinule 

0-9-1 -Omm 
1-0-1-1 mm 
noticeably curved 
0-9-1 -Omm 
3 + 1 setae 

1-1-1-2 mm 
1-1-1-2 mm 
(not exceeding C.L.) 

not pronounced 
average of 8 setae 
average of 1 7 setae 

longer than rostrata 
3 + 1 setae 

average 1*5 mm 
continuous median carina 

forming wide U-shape 

with large spine and 
secondary posterior 

Lebour (1928) provided a key to the larvae of five oxyrhynch genera that occur in British 
waters; Bourdillon-Casanova (1960) extended this number to seven genera in her key to 
Mediterranean brachyuran larvae. From a study of reared material and of published 
accounts (see footnote to Table 2), it has been possible to tabulate a sufficient number of 
comparative larval features from which a provisional key has been constructed to facilitate 
the identification of larvae of all those oxyrhynch genera represented in the North East 
Atlantic Ocean north of about 48 30' N. 

Detailed studies of larval stages of Achaeus cranchii, Pisa tetraodon, Maja squinado, 
Macropodia linaresi and M. dejlexa are still required whilst the positive identity (see Rice, 
1980 : 308) of the plankton caught zoea II attributed to Dorynchus thomsoni still remains to 
be established. 




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Abdomen posterio-lateral spines: 

2 setae on 1st abdominal segment: 

Setae on 2nd abdominal segment: 

Dorso-lateral proc. on 3rd abd. sgm 
Setal formula basis 1st maxilliped: 

No. setae basis of 2nd maxilliped: 

Setal formula endopod 2nd maxillip 

Setal formula endopod maxillule: 
Telson dorsal spine + or -: 


No. abdominal segments (incl. telso 
No. pairs of pleopods (incl. uropods 
No. setae on each terminal pleopod: 

Frontal region of carapace-submedi 

Carapace, each protogastric region i 

orbital region with: 

cardiac region with: 
Spines on abdominal segments: 
Processes or spine, 1 st ped. sgmt anl 

c u 


Provisional key to the larvae of N.E. Atlantic Oxyrhyncha 


Carapace lateral spines present 2 

Carapace lateral spines absent 5 

Carapace with a group of 5 lateral spines on each side 

t Dory nchusthomsoni Thomson, (Williamson, 1960). 

Carapace with one lateral spine on each side 3 

Telson medio-lateral margin strongly concave or medially cleft; middle pair of medio-lateral 
setae longest; each telson furca with one dorsal and one (rarely 2) lateral spines; 3rd 
abdominal segment with dorso-lateral process 4 

Telson medio-lateral margin not concave or cleft; medio-lateral setae of equal length; each 
telson furca without a dorsal spine, one lateral spine present; 3rd abdominal segment 
without dorso-lateral process . . . . Rochinia carpenteriThomson, (Ingle, 1979). 

Carapace dorsal spine much longer than carapace length; abdominal posterio-lateral process 
very long; 2nd maxilliped basis with 4 setae; each furca of telson with one lateral spine 
Hyas coarctatus (Leach), H. araneus (Linnaeus), (Christiansen, 1973). 

Carapace dorsal spine, at the most, slightly longer than carapace length; abdominal 
posterio-lateral processes short; 2nd maxilliped basis with 3 setae; each furca of telson 
with 2 lateral spines Majasquinado(Herbst),(Lebour, 1927, 1928). 

Carapace rostral spine long; each telson furca with a dorsal spine 

Eurynome aspera (Pennant), (Lebour, 1 928). E. spinosa Hailstone, (see Table 2, footnote). 

Carapace rostral spine small or absent; each telson furca without a dorsal spine . . 6 

Telson medio-lateral margin strongly cleft, marginal setae in 2 groups; maxillule endopod* 

with + 3-4 setae; abdominal posterio-lateral spines long 7 

Telson medio-lateral margin not cleft, marginal setae not in 2 groups; maxillule endopod 

with 1 + 6 setae; abdominal posterio-lateral spines short 

tPfaa annaia(Latreille), (Ingle & Clark, 1980). 

Lateral spine of each telson furca very long (about \ length of furca and exceeding ^ maximum 

width of telson) Achaeus cranchii Leach, (Bocquet, 1954). 

Lateral spine of each telson furca short, never reaching | length of furca .... 8 

Lateral spine of each telson furca large; middle medio-lateral seta of telson not longest; 
maxillule endopod with + 4 setae 

Inachus dorsettensis (Pennant), /. p/ialangium (Fabricius), /. leptochirus Leach, (Clark, 1980). 
Lateral spine of each telson furca small; middle medio-lateral seta of telson longest; 
maxillule endopod with + 3 setae 

Macropodia rostrata (Linnaeus), M . tenuirostris (Leach), (see p. 2 1 1 ), M . deflexa Forest (Lebour, 



Abdomen composed of 5 segments (excluding telson); 4 pairs of pleopods, terminal pair with 

8 setae on exopod 2 

Abdomen composed of 6 segments (excluding telson); 5 pairs of pleopods, terminal pair with 

3-5 setae on exopod 4 

Dorsal margins of abdominal segments without spinules; carapace submedian spines not 


Macropodia rostrata (Linnaeus), M. tenuirostris (Leach) (see p. 2 1 1 ), M. deflexa Forest (Lebour, 

Dorsal margins of abdominal segments with small spinules; carapace submedian spines well 

developed 3 

Orbital spines present 

Inachus dorsettensis (Pennant), /. phalangium (Fabricius), /. leptochirus Leach, (Clark, 1 980). 
Orbital spines absent Achaeus cranchii Leach, (Bocquet, 1954). 

*Not known for Achaeus cranchii; tHeegaard's (1963) description of zoea I of Pisa tetraodon differs considerably 
from that of P. armata (see Table 2, footnote) and is excluded from this present key. 

214 R. W. INGLE 

4 Carapace submedian spines present 

Hyas coarctatus (Leach), H. araneus (Linnaeus), (Christiansen, 1973). 

- Carapace submedian spines absent 5 

5 First peduncular segment of antenna without a distal process; exopod of each terminal 

pleopod with 3 setae 

Eurynome aspera (Pennant), (Lebour, 1928), E. spinosa Hailstone, (see Table 2, footnote). 
First pedunular segment of antenna with an obtuse process or spine; exopod of each 
terminal pleopod with 5 setae 6 

6 A stout cardiac spine on carapace . . . Rochinia carpenteri (Thomson), (Ingle, 1979). 
Without a cardiac spine on carapace 7 

7 Rostral spine well developed; each protogastric region with a carina; exopods of lst-4th 

pereiopods with 10 setae Pisa armata (Latreille), (Ingle & Clark, 1 980). 

- Rostral spine minute; each protogastric region with (at the most) a tubercle; exopods of 

1 st-4th pereiopods with 8 setae . . . A/a/a squinado (Herbst), (Lebour, 1 927, 1 928). 


I wish to thank Dr R. B. Manning for sponsoring my visit to Tunisia under the Smithsonian 
Institution Foreign Currency Program; Dr D. I. Williamson for his assistance in obtaining 
ovigerous crabs from the Isle of Man region, and Mr A. D. M. Mattacola and crew of the 
Sepia for their help in collecting material in Plymouth waters. 


Barnard, K. H. 1950. Descriptive Catalogue of South African Decapod Crustacea (Crabs and Shrimps). 

Ann. S. Afr. Mus. 38 : 837 pp. 
Bocquet, C. 1954. Developpment larvaire d'Achaeus cranchii Leach (Decapode Oxyrhynque). Bull. 

Soc. zool. Fr. 79 : 50-56. 
Bourdillon-Casanova, L. 1960. Le meroplancton du Golfe de Marseille: les larves de crustaces 

decapodes. Red Trav. Stn mar. Endoume3Q : 1-286. 

Cano, G. 1 893. Sviluppo e Morfologia degli Oxyrhynchi. Mitt. zool. Stn Neapel 10 : 527-583. 
Christiansen, M. E. 1969. Marine invertebrates of Scandinavia, No. 2. Crustacea Decapoda Brachyura. 

Universitetsforlaget, Oslo, 1-143 pp. 
1973. The complete larval development of Hyas araneus (Linnaeus) and Hyas coarctactus (Leach 

(Decapoda, Brachyura, Majidae) reared in the laboratory. Norw. J. Zool. 21 : 63-89. 
Clark, P. F. 1980. British spider crabs of the genus Inaehus; a morphological study of larval 

development. M.Sc Modern Taxonomy (C.N.A.A.) Thesis. Polytechnic of Central London, 140 pp. 

Heegaard, P 1963. Decapod larvae from the Gulf of Napoli hatched in captivity. Vidensk. Meddr 

dansk naturh. Foren. 125 : 449^493. 
Ingle, R. W. 1979. The larval development of the spider crab Rochinia carpenteri (Thomson) 

[Oxyrhyncha: Majidae] with a review of majid subfamilial larval features. Bull. Br. Mus. nat. Hist. 

(Zool.) 37: 47-66. 

& Clark, P. F. 1 977. A laboratory module for rearing crab larvae. Crustaceana 32 : 220-222. 

1980. The larval and post-larval development of Gibb's spider crab. Pisa armata (Latreille) 

[family Majidae: subfamily Pisinae], reared in the laboratory. /. nat. Hist. 14 : 723-735. 
Ingle, R. W. & Manning, R. B. (in press). Infraspecific variation, synonomy and distribution of the 

spider crab, Macropodia rostrata (Linnaeus). Colloquium Crustacea Decapoda Mediterranean May 

1979 Ancona. 
Lebour, M. V. 1927. Studies of the Plymouth Brachyura. I. The rearing of crabs in captivity, with a 

description of the larval stages of Inaehus dorsettensis. Macropodia longirostris and Maia squinado. 

J. mar. biol. Ass. U.K. 14 : 795-820. 

1928. The larval stages of the Plymouth Brachyura. Proc. zool. Soc. Lond. 2 : 473-560. 

Lo Bianco, S. 1904. Pelagische Tiefseefischeri der "Maja" in der umgebung von Capri. Beitr. Kennt. 

Meeres und Seiner Bewohner 1 : 1-91 Jena. 
Manning, R. B. & Holthius, L. B. 1981. West African Brachyuran Crabs (Crustacea: Decapoda). 

Smithson. Contr. Zool. 306 : 1-379. 


Monod, T. 1932. Brachyuran Maroccana 2e Partie: Dromiidea, Oxystomata, Oxyrhyncha, 

Brachyrhyncha(excl. Pinnotheridae). Bull. Soc. Sci. nat. Maroc 12 : 7-8. 
Rice, A. L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Trans. 

zool. Soc. Lond. 35 : 271^424. 
& Ingle, R. W. 1975. The larval development of Carcinus maenas (L.) and C. mediterraneus 

Czerniavsky (Crustacea, Brachyura, Portunidae) reared in the laboratory. Bull. Br. Mus. nat. Hist. 

(Zool.) 28: 101-119. 
Steedman, H. F. 1976. General and applied data on formaldehyde fixation and preservation of marine 

zookplankton (pp. 103-171) In: Steedman, H. F. (Edit.). Zooplankton fixation and preservation. 

Monographs on oceanographic methodology 4 : 350 pp. UNESCO Press, Paris. 
Stuxberg, A. 1874. Karcinologiska iaktagelser. Ofvers. K. VetenskAkad. Forh. Stockh. 9 : 3-23. 
Thompson, J. V. 1836. Of the double metamorphosis in Macropodia phalangium, or spider crab, with 

proofs of the larvae being zoea in Gegarcinus hydrodomus. Thelpusa erythropus. Eriphia carribaea. 

and Grapsus pelagicus. Ent. Mag. 3 : 370-375. 
Williamson, D. I. 1960. A remarkable zoea, attributed to the Majidae (Decapoda, Brachyura). Ann. 

Mag. nat. Hist. (13)3 : 141-144. 
Zariquiey Alvarez, R. 1968. Crustaceos Decapodos Ibericos. Investigacion pesq. 32 : i-ix + 5 10 pp. 

Manuscript accepted for publication 3 July 1981 



Fig. 1 Macropodia rostrata (L.): a, b zoea I & II respectively from lateral aspects; c zoea II 
carapace and eyes from frontal aspect; d megalopa from dorsal aspect and e lateral aspect of 
carapace and eye; f dorsal and g lateral aspect of megalopal abdomen; h first and i fourth left 
megalopal pleopods; carapace of the first crab from j-Isle of man cf. with k-Tunisian waters. 


1 002' 

Fig. 2 Macropodia rostrata (L.): zoea I, a antennule and antenna, b mandible; zoea II, c 
antennule and antenna, d mandible; megalopa, e antennule, f antenna, g mandible; first crab, h 
antennule & antenna, i mandible. 



Fig. 3 Macropodia rostrata (L.): maxillule of- a zoea I; b zoea II; c megalopa; d first crab. 


Fig. 4 Macropodia rostrata (L.): Maxilla of- a zoea I; b zoea II; c megalopa; d first crab. 



Fig. 5 Macropodia rostrata (L.): a & b 1st & 2nd maxillipeds of zoea I; c & d 1st & 2nd 

maxillipeds of zoea II. 


Fig. 6 Macropodia rostrata (L.): abdominal segments and telson a & c lateral, b & d dorsal 
aspects of- a, b zoea I; c, d zoea II; e spines on right medio-lateral margin of telson of zoea I. 



Fig. 7 Macropodia rostrata (L.): 1 st (a, c) & 2nd (b, d) maxillipeds a, b megalopa; c, d first crab. 


Fig. 8 Macropodia rostrata (L.): 3rd maxilliped- a megalopa; b first crab; cheliped- c megalopa; 

d first crab. 



Fig. 9 Macropodia rostrata (L.): a 2nd pereiopod of megalopa; b, c 2nd & 3rd pereiopods of 1st 



Fig. 10 Macropodia rostrata (L.): a-c 3rd-5th pereiopods of megalopa; d & e 4th & 5th 

pereiopods of first crab. 

British Museum (Natural History) 
British Marine Amphipoda: Gammaridea 

by R. J. Lincoln 

658pp 2,300 figures 4to hard bound 
ISBN 565 00818 50.00 

Amphipods are both numerous and diverse in numbers of genera and species in 
British coastal waters, but in the absence of any form of modern systematic synopsis 
or key this group of crustaceans has acquired the reputation of being notoriously 
difficult to identify. This monograph, which is the first comprehensive and illustrated 
text on British gammaridean amphipods to be published in more than a century, 
should go a long way towards solving the problem. 

The systematic section of the book contains descriptions and figures of all 271 species 
of marine and brackish water amphipods, in 123 genera and 36 families, recorded 
from British coasts and the adjacent continental shelf to a depth of 200 metres. Keys 
are provided at all levels, as well as relevant synonymies and diagnoses of genera and 
families. The text is illustrated with about 2,300 separate figures which have been 
drawn by the author from Museum and other material, in many cases with reference 
to type specimens. 

The work has been carefully edited to bring corresponding descriptions, keys and 
figures into close proximity within the text. The systematic section is supported by 
chapters dealing with morphology, systematics, geographical distribution, biology 
and ecology, the latter being presented in the form of an annotated subject index of 
research literature. Finally, there is an extensive bibliography of about 1,200 
references that includes most of the British marine amphipod literature published to 

Publications Sales, 

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Titles to be published in Volume 42 

The tick collection (Acarina: Ixodoidea) of 

the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and 

general collections of the British Museum (Natural History). 

By James E. Kekans 

Hydroids and medusae of the family Campanulariidae recorded 
from the eastern North Atlantic, with a world synopsis of genera. 
By P. F. S. Cornelius 


Printed by Henry Ling Ltd, Dorchester 


British Museum (Natural History) 


Zoology series Vol 42 No 4 24 June 1982 

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Zool.) 

Trustees of the British Museum (Natural History), 1982 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology : Dr J. G. Sheals 
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Assistant Editor : Mr"C. G. Ogden 

ISSN 0007-1498 Zoology series 

Vol 42 No 4 pp 227-335 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 24 June 1982 



a* ; 



New species of marine nematodes from Loch Ewe, Scotland. By H. M. Platt and 
Z.N.Zhang .............. 

The larval development of Crangon crangon (Fabr. 1795) (Crustacea: Decapoda). 
By A. R.Gurney ............ 

A revision of the spider genus Cocalodes with a description of a new related genus 
(Araneae: Salticidae). By F. R. Wanless ........ 

Anatomy and evolution of the jaws in the semiplotine carps with a review 
of the Genus Cyprinion Heckel, 1 843 (Teleostei: Cyprinidae). By G. J. Howes 


A*^<?ittl ir.AT/nM^"O^ 

New species of marine nematodes from Loch Ewe, 

H. M. Platt 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Z. N. Zhang 

Shandong College of Oceanography, Qingdao, Shandong Province, People's Republic of 
China 1 


During an investigation of the shallow sublittoral nematode fauna off a small sandy bay in 
Scotland one of us (Z.N.Z.) encountered a number of previously undescribed species. These 
were studied in detail at the British Museum (Natural History) and we here present 
descriptions of six of the more interesting species: Gerlachius novusetosus, Catanema 
macintyrei, Catanema smo, Ceramonema yunfengi, Acantholaimus ewensis and Rhips 
paraornata. A new diagnosis is given for the subfamily Gerlachinae Andrassy, 1976. The 
genus Robbea Gerlach, 1956 is synonymized with Catanema Cobb, 1920; a new generic 
diagnosis and a key to the species is given. Keys are also provided for the genera 
Ceramonema Cobb, 1920 and Acantholaimus Allgen, 1933. Several other minor 
nomenclatorial changes are suggested. Species belonging to these five genera have not been 
found previously in British waters and Rhips Cobb, 1920 has not previously been recorded 
from Europe. 

Material and methods 

All specimens came from the same place; sublittoral sand in 3 m of water off 'north beach', 
Firemore Bay, Loch Ewe, Wester Ross, Scotland. Detailed descriptions of the bay and its 
fauna can be found in Steele & Baird (1968), Mclntyre & Eleftheriou (1968) and Mclntyre & 
Murison (1973). The glycerine mounted specimens were studied using a Leitz Ortholux II 
microscope equipped with differential interference contrast facility. All drawings were made 
using a drawing tube. The formulae used for the dimensions are a modification of Filipjev's 
(1918) formula and the de Man ratios, as described in Platt (1973). The abbreviations 'S' and 
'V are the spicule length and the relative position of the vulva respectively. The 
abbreviations 'a.b.d.' and 'c.d.' are the anal (or cloacal) body diameter and corresponding 
(body) diameter. Type material has been deposited at the British Museum (Natural History). 

Systematic descriptions 

The classification follows that given in Gerlach & Riemann (1973). 

Subfamily GERLACHINAE Andrassy, 1976 

DIAGNOSIS. Meyliidae. Cuticle smooth or feintly striated. Amphids circular, lightly 
cuticularised or invisible. Testes paired, opposed. Ovaries reflexed. 

'also, D.A.F.S. Marine Laboratory, Victoria Road, Aberdeen AB9 8DB 

Bull. Br. Mus. nat. Hist. (Zool.) 42 (4) : 227-246 Issued 24 June 1 982 



DISCUSSION. Andrassy (1976) erected the genus Gerlachius to distinguish Meylia lissa 
Gerlach, 1956 from the other two species of the genus, M. alata Gerlach, 1956 and 
M. spinosa Gerlach, 1956. According to Andrassy, Gerlachius is characterized by its 
smooth cuticle and invisible amphids: the specimens described below are similar to 
G. lissus in many respects apart from these two main generic characters. However, 
the cuticle in G. novusetosus sp. nov. was observed under interference contrast and 
can otherwise be seen only as feintly striated so that this feature may have been over- 
looked in G. lissus. Likewise, the amphid may have been overlooked, since in the 
male G. novusetosus sp. nov. it was difficult to distinguish. Therefore, at this stage we 
prefer simply to widen the subfamily and genus diagnosis to permit forms with a smooth 
cuticle without amphids until these animals become better known. 

The subfamily Gerlachinae can now be distinguished from Meyliinae since the ovaries are 
outstretched in the latter (Lorenzen, 198 1) but reflexed in the former. 

Fig. 1 Gerlachius novusetosus: (a) whole body of d; (b) head of rf; (c) head of 9; (d) copulatory 
apparatus of rf; (e) 9 reproductive system. Bar scales: a,e = 40 um; b,c,d = 10 um. 


Gerlachius novusetosus sp. nov. 
Fig. 1 

MATERIAL STUDIED. Holotype: d 1 BM(NH) 1981.4.10. 
Allotype: 9 BM(NH) 1 98 1 .4. 1 1 . 


Holotype rf: - 34 M 433 505 /zm;a = 23;b= 14-9; c = 7-0; S = 31 um 

11 18 22 17 

Allotype 9: - 34 280 463 515/zm;a = 19;b= 15-l;c = 9-9; V = 54% 

12 22 27 17 

DESCRIPTION. Short, relatively stout body. Cuticle feintly striated. Short 2'5-3-5//m 
sublateral somatic setae present (Fig. la): they are not bilaterally symmetrical. Four 5*5 um 
submedian cephalic setae (R3?) seated on short l - 5 um peduncles. A short stout additional 
seta, 2-2-5 um long, is associated with each cephalic seta positioned as shown in Fig. Ib, c. 
Amphid round, not strongly cuticularised, 7 um long and 6*5 um wide in female (male 
amphid less distinct but about 5-5 jum long). Posterior to the amphid there is a subcuticular 
reticulate structure (Fig. Ic) of unknown function. Buccal cavity absent. Oesophagus short, 
without a bulb. Tail conical, 3-4 a.b.d. Three conspicuous caudal glands lying entirely in the 

Spicules curved and slightly cephalate proximally: chord length 23 um, arc length 3 1 um. 
Gubernaculum has a dorso-caudally directed apophysis. Anterior to cloaca the cuticle 
striations are thickened ventrally to produce a longitudinal row of contiguous pegs which 
extends about 190 um from the cloaca (Fig la, d). Testes paired, opposed. 

Vulva cuticularised and prominent. Ovaries paired, opposed, reflexed. 

DIFFERENTIAL DIAGNOSIS. Gerlachius novusetosus sp. nov. differs from the only other 
species, G. lissus (Gerlach, 1956) in having four short additional cephalic setae, shorter 
cephalic setae (5*5 um vs 15 um), shorter oesophagus (b = 15 vs 7) and precloacal cuticular 

DISCUSSION. The reticulate organ posterior to the amphid and the four additional pegs 
alongside the cephalic setae have not been reported previously. However, this whole group 
of what appear to be primitive desmoscolids are poorly known, the only other records being 
G. lissus (Gerlach, 1956), Meylia alata Gerlach, 1956 and M. spinosa Gerlach, 1956; all 
from Kiel Bay. 

CATANEMA Cobb, 1920 

Robbea Gerlach, 1956 syn. nov. 

The genus Catanema was originally described by Cobb, 1920 to accommodate C. exile from 
Jamaica, a male specimen with fine cuticle striations, narrow buccal cavity 'enclosed in a 
swelling', seven pairs of subventral postcloacal tubular organs, proximally cephalate spicules 
and a large dorso-caudally directed gubernacular apophysis. Three further species have 
subsequently been assigned to the genus: C. cobbi Inglis, 1968, with ten pairs of subventral 
postcloacal organs, cephalic cuticle 'modified into blocks' and a dorsally directed 
gubernacular apophysis; C. porosum Hopper & Cefalu, 1973, with five pairs of subventral 
postcloacal organs; C. gerlachi Hopper & Cefalu, 1973. This last species was a new name 
for a specimen from the Maldive Islands originally described by Gerlach (1963#) under the 
name Eubostrichus exilis (Cobb, 1920). However, as Hopper & Cefalu (1973) point out, 
Gerlach (19630) makes no mention of the buccal bulb so characteristic of this genus and the 
gubernaculum lacks the prominent apophysis, which in Gerlach's specimen is thin and lies 
almost parallel to the spicules. 


Hopper & Cefalu (1973) decided to keep this record within the genus Catanema. 
However, we feel that it is more probable that Gerlach would not have overlooked a buccal 
bulb if it were present (having already described nematodes with this character, e.g. Robbea 
caelestis Gerlach, 1956) so we transfer this species back to Eubostrichus, becoming 
Eubostrichus gerlachi (Hopper & Cephalu, 1973) comb. nov. Eubostrichus is characterized 
as having modified 'ponds' on the tail (Hopper & Cefalu, 1973: porids = tubular setae serving 
as outlets for glands): Gerlach (1963 p. 95) depicts similar structures in both E. parasitiferus 
Chitwood, 1936 and his E. exists ( = E. gerlachi) from the Maldive Islands and both species 
have similarly shaped guberaacula. 

In 1956, Gerlach erected the genus Robbea for a male specimen from Brazil, considering 
that the muscular buccal bulb distinguished the taxon from all other related genera. 
Although the type, R. caelestis, is certainly depicted as having a very prominent buccal bulb, 
some species described subsequently seem to have somewhat less prominent bulbs, depicted 
as similar to those described for Catanema. This being so, we propose to synonomize 
Robbea with Catanema so that Catanema now also includes the following species: C. 
caelestis (Gerlach, 1956), C. gallica (Vitiello, 1974) and C. tenax (Gerlach, 19636). C. 
gerlachi (Boucher, 1975) was described from a female only: since male characters are of 
importance in this taxon, and despite the fact that C. gerlachi seems to be unique in the 
length of its subcephalic setae, we prefer to treat it as a species inquirenda. 

In proposing this synonomy, there remains two main points to discuss: the amphid of the 
type and the presence of subventral tubular supplements on the male tail. In Cobb's (1920) 
description of the type, C. exile, he describes the amphids as 'minute labial tubes'. However, 
he also states that they were 'forward-pointing . . . difficult to see'. A similar anterior position 
was found in C. smo sp. nov. described here and we feel confident that Cobb misinterpreted 
what were spiral, albeit minute, amphids. Cobb (1920) also depicted the type with seven 
prominent pairs of subventral tubular supplements on the tail, similar to the modified porids 
found in Eubostrichus. We cannot be sure whether these supplements, found in C. exile, C. 
cobbi and C. porosum are homologous with the stout caudal setae described in C. caelestis, 
C. gallica, C. tenax and the two new species described here. However, their presence or 
absence seems to be no more significant a reason for suggesting a generic split than say the 
presence of huge cervical suckers in C. tenax or the reticulate head of C. cobbi. Therefore, we 
consider the presence or absence of caudal supplements as a variable infrageneric character. 

Finally, we would comment on the arrangement of setae on the head. Inglis (1968) 
described 6 + 4 cephalic setae, the lateral components of the R 2 circle being 'markedly dorsal' 
in position (although, according to the direction of the amphid spiral, he mistook ventral for 
dorsal). However, in most species, there are four prominent setae, often longer than the 
others; we judge these to be the four R 3 sensilla. The six R 2 sensilla are small in size, often not 
depicted but can be clearly seen in the two species described here. If so, the R, sensilla must 
be reduced to papillae and may require SEM to resolve. In all species, each R 3 sensillum 
seems to be accompanied by a more medially situated subcephalic seta, which may be at 
about the same level as the R 3 sensilla as in C. tenax or more posterior as in C. caelestis. It is 
these which Inglis (1968) called the four cephalic setae. In addition to these subcephalic 
setae, there are others on the cephalic capsule which, according to the several specimens of 
each species described here, seem to be quite stable in position. Most species, including C. 
smo sp. nov., have setae located either side of the base of the amphid. In at least C. cobbi, C. 
macintyrei sp. nov. and C. tenax there are additional setae in specific locations. For example, 
in C. cobbi there is a pair of setae either side of the base of the amphid and an additional seta 
associated with each ventral subcephalic seta. In C. macintyrei sp. nov. there is a seta close 
behind each subcephalic seta, a small seta associated with each ventral R 3 seta and a seta 
lying dorsal to the amphid. 

We propose the following generic diagnosis: Catanema. Desmodoridae. Stilbonematinae. 
Elongated body. Cuticle finely striated posterior to cephalic capsule and in life covered with 
coccoid blue-green algae. Cephalic capsule may appear reticulate or feintly punctated. Four 
anteriorly directed R 3 cephalic setae and four subcephalic setae. Additional subcephalic 



setae may be present on the cephalic capsule. Spiral amphid. Buccal cavity minute and 
funnel-shaped leading to a muscular buccal bulb. Oesophagus narrow, terminating in a 
rounded posterior bulb. Spicules curved and proximally cephalate. Gubernaculum with a 
solid dorsally or dorso-caudally directed apophysis. Single outstretched testis. Subventral 
pairs of tubular organs may be present on the tail. Tail conical. 

TYPE SPECIES. Catanema exile Cobb, 1920 

Key to the species of Catanema 

1 Ventral suckers present in oesophageal and post-oesophageal region 

C. tenax (Gerlach, 1963) comb. nov. 

syn. Robbea tenax Gerlach, 1963 

Ventral suckers absent 2 

2 Cephalic cuticle in conspicuous blocks. 1 pairs subventral caudal supplements 

C.coAA/Inglis, 1968 
- Cephalic cuticle smooth or with feint punctations 3 

3 Tail tip pointed, 'a' ratio > 200. Amphid large (0-7 c.d.) C. caelestis (Gerlach, 1 956) comb. nov. 

syn. Robbea caelestis Gerlach, 1956 
Tail tip rounded, 'a' ratio < 200 4 

4 Amphid situated far anterior so that subcephalic and subamphidal setae are on the same 

level. 7 pairs subventral caudal supplements C. exile (Cobb, 1920) 

Amphid situated further posterior 5 

5 Middle of amphid situated at 10% of length of buccal bulb from anterior . . C. smosp. nov. 
Middle of amphid situated at least 30% of length of buccal bulb from anterior . . . 6 

6 Cephalic setae 8 //m long (0-3 c.d. at base of buccal bulb) C. gallica (Vitiello, 1 974) comb. nov. 

syn. Robbea gallica Vitiello, 1974 
Cephalic setae > 17 ^m long (0-8 c.d. at base of buccal bulb) 7 

7 Only 4 subcephalic setae and 4 subamphidal setae between base of amphid and anterior 

C. porosum Hooper & Cefalu, 1973 
Additional subcephalic setae present C. macintyrei sp. nov. 

Catanema macintyrei sp. nov. 
Figs 2-3 

MATERIAL STUDIED. Holotype: rf, BM(NH) 1981.5.22. 
Allotype: 9, BM(NH) 1981.5.23. Paratypes: five 


males and five females BM(NH) 

Allotype 9,: 

- 87 M 3710 3770//m;a= 

25 25 27 23 
- 94 1845 3555 3620 //m; a = 125;b = 39;c = 56; V = 51% 

25 25 29 24 

= 44//m 

a=138; b = 37; c = 66; 

a=137; b = 42; c = 71; 

a=143; b = 47; c = 72; 

a=146; b = 45; c = 64; S = 43/zm 

a=128; b = 41; c = 61; S = 41/an 

9 2 :L = 4065/*m; a=145; b = 47; c = 71; V = 50% 

9 3 :L = 3720/zm; a =143; b = 40; c = 69; V = 50% 

9 4 :L = 4510//m; a=161; b = 46; c = 76; V = 52% 

9 5 :L = 4010/zm; a =143; b = 47; c = 78; V = 52% 

9 6 :L = 4530//m; a =162; b = 49; c = 79; V = 49% 



Fig. 2 Catanema macintyrei: (a) whole body of dy, (b) oesophageal region of cf,; (c) head of cf, (see 
text for labelling); (d) head of d" 7 ; (e) head of 9,. Bar scales: a = 100 //m; b = 30 ^m; c-e = 10 ^m. 

Spicule (S) measurement is the arc. Maximum body diameter used is the average of three 
measurements along the middle portion of the body. 

DESCRIPTION. Cuticle finely striated from the base of the cephalic capsule, but most 
conspicuous in the oesophageal and immediate post-oesophageal regions. Typically, the 
cuticle is covered with a coating of coccoid blue-green algae, but this tends to be lost during 
preservation. One specimen also had two suctorian ectoparasites attached to the middle part 



Fig. 3 Catanema macintyrei: (a) tail region of c? 2 ; (b) copulatory apparatus of d 1 ,; (c) spicules of 
three different males; (d) tails of three males and two females. Bar scales: a,c = 30 jum; b = 10 /zm; 

of the body. Cuticle bears six files of stout setae throughout the body, but they are 
particularly conspicuous ventrally in the region just posterior to the oesophagus and in the 
male, anterior to the cloaca. R, sensilla not seen. Six small l'5-2 /zm setose R 2 sensilla. Four 
17'5-24 //m R 3 setae, about 80% of the maximum diameter of the cephalic capsule. The head 
diameter is usually measured at the level of the R 3 sensillae, but they are so far anterior as to 
make this measurement too inaccurate. Medial and slightly posterior to the R 3 setae are four 
ll-14//m subcephalic setae (labelled SC, in Fig. 2c). Further posterior are four smaller 
6-5-8*5 jum setae (labelled SC 2 in Fig. 2c). In addition, there is one seta just posterior to the 
ventral R 3 seta (Fig. 2c, V), two level with and dorsal to the middle of the amphid (Fig. 2c, D, 
& D 2 ) and one ventro-sublateral subamphidal seta (Fig. 2c, A). The disposition of these setae 
is constant in that each element can be distinguished in the same location in all the 
specimens, both male and female. Amphid ventrally wound, from outside to centre; 
9-10//m wide. Buccal cavity represented by a narrow funnel in the anterior part of the 
buccal bulb, the latter being about 23 //m long. Tail conical with a characteristic slight 
ventral inflection at the tip. 

Spicules paired, curved and proximally cephalate: average length 44 jam (arc) or 32 fj.m 
(chord). Gubernaculum well cuticularised and dorso-caudally directed. Testis single and 

Ovaries paired and apparently outstretched. 

DIFFERENTIAL DIAGNOSIS. Catanema macintyrei sp. nov. can be distinguished from the other 
species in the genus Catanema by the unique disposition of the setae around the head. From 


the other sympatric species, C. smo sp. nov., it can also be distinguished by the larger and 
slightly more posteriorly situated amphid. 

DISCUSSION. This species will be discussed together with the following species. 

Catanema smo sp. nov. 
Fig. 4 

Fig. 4 Catanema smo: (a) oesophageal region of cf,; (b) head of cf,; (c) head ofd 1 ,; (d) tail region of 
rf 2 ; (e) copulatory apparatus of cf,; (0 spicules of three different males; (g) tails of three males and 
one female. Bar scales: a,d,f = 30 //m; b,c,e = 1 //m; g = 50 //m. 


MATERIAL STUDIED. Holotypeicf, BM(NH) 1981.5.38. 

Allotype: 9, BM(NH) 1981.5.39. Paratypes: four males BM(NH) 1981.5.35-37,30. 


Holotyperf,: -83 M 2980 3050 /on; a = 122;b = 37;c = 43;S = 38/mi 

24 24 25 24 

Allotype 9,: - 80 1900 3393 3460/un;a=121;b=44;c=52; V=55% 

24 25 29 21 

= 3090/mi; a=125; b = 51; c = 54; 

= 3285/mi; a=131; b = 40; c = 54; 

a=110; b = 38; c = 50; 

a=131; b = 38; c = 53; 

(Spicule and maximum body diameter measured as in C. macintyrei). 

DESCRIPTION. Cuticle finely striated from the base of the cephalic capsule, but most 
conspicuous in the oesophageal and immediate post-oesophageal regions. Typically, the 
cuticle is covered with a coating of coccoid blue-green algae, but this tends to be lost during 
preservation. The cuticle of the cephalic capsule has a feint punctated or dotted appearance 
viewed with interference contrast microscopy although this is difficult to depict: it is not 
obvious with ordinary illumination. There are rows of setae in the oesophageal region but 
somatic setae are absent from the rest of the body. R, sensilla not seen. Six small (about 
1 //m) setose R 2 sensilla. Four 17-21 //m R 3 setae, about 80% of maximum cephalic capsule 
diameter and situated at the anterior extremity of the head. Four shorter subcephalic setae 
and four sublateral subamphidal setae. This disposition of the setae on the anterior part of 
the head was the same in all specimens. Amphid ventrally wound, situated just posterior to 
the R ? setae and 6-7 //m wide. From a lateral view, the amphid appears oval but when the 
head is bent slightly the amphid can be seen to be round (Fig. 4c). Buccal cavity a narrow 
funnel in the anterior part of the buccal bulb. Tail conical with a characteristic ventral 
inflection in the slightly more cylindrical terminal portion. 

Spicules paired, curved and proximally cephalate: average length 36 //m (arc) or 27 //m 

(chord). The proximal cephalation has a characteristic knob which is easy to distinguish 

even at low magnification (Fig. 4f). Gubernaculum well cuticularised and dorso-caudally 

^directed. Testis single and outstretched. Tail has three pairs of subventral setae and a single 

precloacal ventral seta was present in at least two males. 

Ovaries paired and apparently outstretched. 

DIFFERENTIAL DIAGNOSIS. Catanema smo sp. nov. can be distinguished from most other 
Catanema species with only four subcephalic and four subamphidal setae by the anterior 
position of the amphid. The only other species with its amphids so far forward is C. exile 
Cobb, 1920, which has 7 well developed pairs of tubular caudal supplements. 

DISCUSSION. Superficially, both the species found in the same samples from Loch Ewe 
appear to be similar. However, on more detailed examination they were shown to be 
separate species on a number of characters. In practice, the more distinct knob at the 
proximal end of the spicule could be used to distinguish males of C. smo at relatively low 
magnification once the existance of two separate species was realized. Other differences 
include the presence of two conspicuous subamphidal setae in C. smo, presence of somatic 
setae throughout the body in C. macintyrei and slightly smaller spicules in C. smo. Both 
species were found mainly 9-12 cm deep in the sand, the epizoic algae being a common 
feature on deeper-living nematodes. 


Ceramonema yunfengi sp. nov. 
Fig. 5(a-Ki) 

MATERIAL STUDIED. Holotype: <? BM(NH) 1 98 1 .4. 1 2. 

Allotype: 9, BM(NH) 1981.4.13. Paratype 9 2 BM(NH) 1981.4.14. 


Holotype cf : - 181 M 1328 1 560 //m; a = 78; b = 8-6; c = 1 1 -7; S = 24 ^m 

16 18 20 18 

Allotype 9,: -125. M ,791 927/rni;a = 54;b = 7-4;c = 6-8; V = 44% 

15 17 17-5. 12 

9 2 : - 141 M ' 874 1020/nn; a = 58;b = 7-2;c = 7-0; V = 46% 

16 17-5 17-5 12-5 

DESCRIPTION. Body colourless, elongated and conspicuously annulated. Cuticle annules in 
male 5'5//m, 7/^m and 5 /*m long in the anterior oesophagus, posterior oesophagus and 
cloacal regions respectively. Total number of body annules 277 in cf, 186 in 9, and 184 in 9 2 . 
Each annule divided into eight plates, so that there are eight longitudinal crests running the 

Fig. 5 Ceramonema yunfengi: (a) whole body of cf,; (b) cf, head; (c) cf, tail; (d) 9, head. Bar scales: 


length of the body. Vacuoles are present beneath the cuticle which are not confined to the 
longitudinal crests. In the male, but not the female, the annule in the cloacal region is 
elongated (about 12//m) relative to the rest (Fig. 5c). Somatic setae absent except for two 
caudal setae situated on the distal cone of the male only. Cephalic capsule elongated: <5 
33-5 x 1 6'5 //m; 9, 29 x 1 5 /zm; 9 2 3 1 -5 x 1 6 //m. Six slim R 2 cephalic setae; 1 2-5 //m long in cf 
and 10-10'5 //m long in 99. Four slim R 3 cephalic setae situated just posterior to R 2 setae; 
13'5 jum long in d, 1 1 //m long in 99. In the d, two medial holes in the cephalic capsule were 
observed just posterior to the level of the R 3 sensilla (Fig. 5b): possibly positions of sub- 
cuticular sensilla. Amphids an elongated loop, 17-1 9 /zm long. Buccal cavity absent. 
Oesophagus narrow but widens slightly at the posterior end. Nerve ring at 52-54% of 
oesophagus length. Excretory ampullae situated anterior and posterior to oesophageal- 
intestinal junction. Tail elongated, 7 a.b.d. in d, 1 1-12 a.b.d. in 99. In the d, the terminal 
cone is 14- 5 //m long and bears two lateral 8 //m setae. 

Spicules paired, slightly curved and 24 //m long (chord). Gubernaculum almost straight, 
1 5 um long. No supplements. Testes not clear. 

Ovaries paired, opposed and reflexed. 

DIFFERENTIAL DIAGNOSIS. Ceramonema yunfengi sp. nov. can be distinguished from all 
other known species of the genus by the presence of two lateral sub-terminal caudal setae in 
the d and a relatively longer cephalic capsule. The only other species described which have 
an elongated cloacal annule are C. carinatum Wieser, 1959 and possibly, according to the 
figure, C. pisanum Gerlach, 1952. However, both these species have shorter cephalic setae 
and are not described as having subcuticular vacuolisation. In addition, the tail of C. 
pisanum is longer than that of C. yunfengi (1 1 a.b.d. vs. 7 a.b.d.). 

DISCUSSION. Haspeslagh (1973) recently revised the superfamily Ceramonematoidea and, on 
the basis of the presence or absence of vacuolisation in the cuticle, divided Ceramonema into 
three genera. However, Lorenzen (1981) did not accept this argument and synonomized 
Haspeslagh's new genera, Ceramonemoides and Cyttaronema, with Ceramonema. 
Currently, there are eleven species contained in the genus including C. yunfengi, which can 
be distinguished using the data contained in Table 1 and Fig. 6. 

Apart from C. pisanum Gerlach, 1952, which is described from a male and a juvenile 
female, C. yunfengi is the only species where both male and adult female characters are 
known. Four species are known from females only and one from a juvenile only. Many of the 
important characters used to distinguish marine nematodes in general are vested in the male, 
so current practice is to consider species described from females or juveniles only as dubious. 
However, it is still possible to separate the eleven species contained in Table 1 on the 
combination of the relative length of the cephalic capsule, presence or absence of 
vacuolisation and its pattern of distribution, relative position of R 2 and R 3 sensilla and 
cuticle structure. However, as more species become known, it may become impossible to 
separate the taxa on these characters alone so that C. attenuatum, C. chitwoodi, C. rectum, C. 
sculpturatum and C. undulatum may have to be designated species dubiae, particularly as 
certain characters display sexual dimorphism including length of R 2 and R 3 sensilla and 
number of body annules, at least in C. yunfengi. The apparent sexual dimorphism in the 
amphid reported for C. pisanum (Fig. 6e, may be simply due to the female not being adult: 
the amphids are similar in C. yunfengi. 

Acantholaimus ewensis sp. nov. 

Figs 7-8 

MATERIAL STUDIED. Holotype: d, BM(NH) 1 98 1 .4. 1 5. 

Allotype: 9, BM(NH) 1981.4.16. Paratypes: d 2 BM(NH) 1981.4.17; d 3 BM(NH) 1981.4.18; 

d 4 , d 5 in collection of Z.N.Z. 



c c c c 



<S23 <^<S 

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D222 D.2D2.0 
8 tt 8 o E tt tt 8 fi 


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<-. | -. | e- e- c- + 

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1 1 +' 1 + 

r* ** 

c B c 

+ 1 1 1 

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^M IO CO *^ OO t 

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.a u 

* * * * 

.C 00 ~!r 

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t^ *O ro *O ^D OJ OO ^" *^ f*^ oo 


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u -> 



(N C?\ t~~ TT OO ~* ^ <^> 


*^ 4> 

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~*Tt<Nm'O(N l /^rOI^<Nt^ 








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I 1 

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Table I Tabular 




_ , (vj r~ ^ S d 2 . rr> "" 

g-^-j in ^ _^ ON ovc""" 1 N a""f'^ 

S^gON.^U X. 3-0' ^ T3 C 

Q t - uT ^ C '5 cS^r'S^ O^ u2S OQ S^> 
s S & $ Q O 9 <B s s 5 S (i ^ 3k l ^ C Q 

^ O Ji '~C 1? V D .^ u 5> U *-i ^ "ts 1> 2 j C " S rv 

^calculated from I 
+'vacuoles arrang 



Fig. 6 Pictorial key to the species of Ceramonema: (a) C. attenuatum 9 (after Cobb, 1920); (b) C. 
carinatum d (after Wieser, 1959); (c) C. chitwoodi 9 (after De Coninck, 1942); (d) C.filipjevi d 
(after De Coninck, 1942); (e) C. pisanum <3 (after Gerlach, 1952); (0 C. pisanum juvenile 9 (after 
Gerlach, 1952); (g) C. rectum 9 (after Gerlach, 1957); (h) C. reticulatum d (after Chitwood, 1936); 
(i) C. salsicum d (after Gerlach, 1956); (j) C. sculpturatum juvenile (after Chitwood, 1936); (k) C. 
undulatum 9 (after De Coninck, 1 942); (1) C. yunfengi sp. nov. 

Holotype rf, : 

Allotype 9, : 

- 115 M 550 
15 23 23 19 

- 125 455 680 930/rni; a = 

17 25 32 23 

= 7-4;c = 3-7; V = 49/o 

cT 2 : L = 860 //m; a = 30; b = 7'5; c = 3'7 

a = 35; b = 7'3; c = 3'5 

a = 29*; b = 7'3; c = 4-2 

a = 25*; b = 7'5; c = 3'5 
4 and d" 5 were slightly flattened. 



Fig. 7 Acantholaimus ewensis: (a) whole body of d 1 ,; (b) head of d 1 ,; (c) head of 9, showing surface 
structures only; (d) head of 9, showing buccal cavity structure. Bar scales: a=100wm; 
b-d= 10//m. 

DESCRIPTION. Cuticle punctated and laterally differentiated. Punctation extends anterior to 
amphids (Fig. 7c) and in the anterior oesophageal region the lateral punctations are irregular. 
Further posteriorly, the punctations become smaller and arranged in transverse rows. In the 
middle of the body, the transverse rows of punctations can be difficult to distinguish. Lateral 
differentiation consisting of two longitudinal rows of larger punctations begins level with the 
middle of the posterior oesophageal bulb and ends at about 65% of the tail length. The lateral 
rows are about 1 -5 /zm apart and there is a space between these rows and the transverse rows 
of punctations (Fig. 8f). There are four sublateral rows of hypodermal pores, which are larger 



anteriorly (Figs 8d-f). Long somatic setae are present throughout the length of the body and 
arranged more or less in four sublateral rows. In the oesophageal region, the length of the 
somatic setae increases from 15-20//m at the anterior to 3 5-40 //m at the posterior. In the 
middle of the body, the longer setae are about 45 //m long but decrease in length again to 
about 30-40 //m in the anal region and 10-20 //m in the tail. 

At the anterior extremity there is a circle of six 2*5-3 /zm stout cephalic sensilla, 
presumably the R 2 sensilla. Just behind are four 30-33 //m R 3 cephalic setae which, in some 
specimens, appear to have a joint at about two-thirds of the length (Fig. 7b). Amphids 
8'5-9'5//m wide (45-50% c.d.), circular with a well cuticularised boarder and a ventrally 
directed posterior inflection. 

Buccal cavity rather complex but basically it is conical with a distinct dorsal and two 
subventral teeth and anteriorly bears twelve rugae. Oesophagus has characteristic plasmatic 
interruptions (Fig. 7d) throughout its length and posteriorly there is a distinct bulb, 
28 x 18 //m. Tail elongated (11-12 a.b.d.), gradually tapering but with a slightly bulbous tip 
and a pointed spinneret (Fig. 8c). 

Fig. 8 Acantholaimus ewensis: (a) tail of d 1 ,; (b) copulatory apparatus of d,; (c) tail tip of $,; (d) d, 
cuticle ornamentation about 40 //m from anterior; (e) d, cuticle ornamentation level with the 
posterior of the oesophagus; (f) d, cuticle ornamentation at middle of body. Bar scales: 
a = 30 //m; b-f = 1 //m. 





Fig. 9 /J/zips paraornata: (a) anterior region of d 1 ,; (b) head of d 1 ,; (c) head of another cf; (d) tail of 
d 1 ,; (e) copulatory apparatus of d 1 ,; (fHO cuticle patterns at positions indicated in Fig. 10. Bar 
scales: a,d = 30 ^m; b,c,e-l = 10 /am. 

Spicules paired, equal, 26-28 ^m long and with a characteristic ventral apophysis. 

Gubernaculum absent. In d 1 ,, a feint cuticularised structure was observed lateral to the 
spicule (Fig. 8b): it was not detected in the other specimens. There is a single stout ventral 
precloacal seta, 5-6 //m long. Testis single and mature specimens contain large pear-shaped 

Ovary paired, opposed and reflexed. In the female studied, there were sperm present in the 
oviduct and a large egg in each uterus containing a developing larva. 

DIFFERENTIAL DIAGNOSIS. Acantholaimus ewensis sp. nov. can be distinguished from 
Acantholaimus species with two lateral longitudinal rows of larger punctations, A. poly- 



Fig. 10 Rhips paraornata: whole body of rf, showing positions of cuticle pattern drawings in Fig. 

9f-l. Bar scale =100 //m. 

dentatus Gerlach, 195 1 and A. calathus Gerlach, Schrage & Riemann, 1979, on the length of 
the R 3 cephalic setae, somatic setae and tail. A. polydentatus has shorter cephalic setae 
(20-22 urn, 1-5 h.d. vs. 30-33 /zm; 2'0 h.d.), shorter somatic setae (30 /zm, 0'75 c.d. vs. 
45 um, 2-0 c.d.) and a shorter tail (8 a.b.d.vs. 11-12 a.b.d.). A calathus has shorter cephalic 
setae ( 1 2 um, 1 -0 h.d.), shorter setae (27 um, 1 -0 c.d.) and longer tail ( 1 7 a.b.d.). 

DISCUSSION. The genus Acantholaimus Allgen, 1933 has been fully revised recently by 
Gerlach, Schrage & Riemann (1979) who described seven new forms and provided a key to 
the twelve known species. A. ewensis fits into the first part of their key as follows: 

1 (4) Cuticle laterally with two longitudinal rows of larger punctations . . 2 

2 (3) Amphid situated close behind the cephalic setae 2 1 

T (2") Tail 8 anal diameters long A. polydentatus Gerlach, 1951. 

1" (2') Tail Tl-12 anal diameters long A ewensis sp. nov. 

3 (2) Amphid situated half the head diameter behind the cephalic seta. Tail 1 7 anal 

diameters long. A. calathus Gerlach, Schrage & Riemann, 1979 

4 (1) Cuticle with limited or no lateral differentiation, without longitudinal rows of 

punctations 5 

Rhips paraornata sp. nov. 
Figs 9- 10 

MATERIAL STUDIED. Holotype: rf, BM(NH) 1 98 1 .4. 19. 

Allotype: 9, BM(NH) 1981.4.20. Paratypes: nine males and two females; rf 6 BM(NH) 

1 98 1 .4.2 1 , remainder in collection of Z.N.Z. 

244 H. M. PLATT & Z. N. ZHANG 


Holotyperf,: - 180 M 990 1 1050m;a = 48;b = 6-l;c = 9-6;S = 7 

11 21 23 22 

Allotypeg,: - 179 M 938 10800m;a = 37;b = 6-0;c = 7-6; V = 54% 

10 26 29 20 

rf 2 :L= 10850m; a = 38; b = 6'l; c=8'7 

dy.L=13300m; a = 50; b = 6'5; c= 9*3 

rf 4 :L=11300m; a = 48; b = 6'5; c=ll'9 

rf 5 :L=12600m; a = 44; b = 6'4; c= 8'5 

d" 6 :L=12600m; a = 47; b = 6*9; c=ll'0 

d" 7 :L= 9200m; a = 42; b = 5'3; c= 8-7 

d- 8 :L=12800m; a = 53; b = 6'7; c=10'0 

cT 9 :L=12750m; a = 53; b = 7'4; c=13'0 

<*w'.L= 11600m; a = 50; b = 6'9; c=9'0 

9 2 :L=10950m; a = 44; b = 6'6; c= 9'0 

9 3 :L= 9800m; a = 36; b = 6'5; c= 7-3 

DESCRIPTION. The cuticle is conspicuously annulated and heterogeneously ornamented 
laterally. In the anterior third of the oesophagus, from about the point where the body 
characteristically narrows (Fig. 9a) each annule bears a row of long and a row of smaller 
round punctations (Fig. 90, the latter being difficult to distinguish and liable to be over- 
looked. In the posterior part of the oesophagus, the punctations are smaller and appear to be 
partly linked diagonally (Fig. 9g), giving a reticulated appearance. The regular nature of the 
ornamentation breaks down posterior to the oesophagus and lateral differentiation of large 
dumb-bell-shaped punctations begins (Fig. 9h). Just posterior to the mid-point of the body, 
the pattern of the lateral differentiation reverses and, at the point of change, the dumb-bell- 
shaped lateral punctation is represented by a single round dot (Fig. 9j). Lateral differentiation 
ends just anterior to the anus and transverse rows of discrete punctations are present on the 
tail (Fig. 91). 

Short 4-6 0m sublateral somatic setae are present at infrequent intervals throughout the 
body and at a third of the oesophagus length, there are four longer somatic setae (Fig. 9a). 
The head bears six setose R, sensilla. The six 2-3 0m R 2 sensilla are situated just anterior to 
the four shorter (1*5-2 0m), R 3 sensilla. The elongated first body annule forms a cephalic 
shield and bears six triangular flap-like extensions anteriorly which alternate with the 
R 2 + R 3 sensilla (Fig. 9b, c). The shield is irregularly punctated and bears the large transverse 
amphids. The amphid has strongly cuticularised borders and is 1 1 0m wide, about 0*85 c.b.d. 
The buccal cavity is surrounded anteriorly by rugae which protrude beyond the lips. There is 
a large pointed dorsal tooth and two smaller subventral teeth. The oesophagus widens 
posteriorly to a weak bulb. The tail is conical and has an unstriated end cone. 

The male copulatory apparatus is complex, consisting of two long double-jointed spicules, 
a paired gubernaculum and two lateral pieces. In the holotype, the posterior part of the 
spicule measures 40 0m as the arc (or 29 0m as the chord) and the anterior part measures 
38 0m as the arc (or 35 0m as the chord). In five other males, the total arc length of the 
spicule is 70-96 0m. The lateral pieces are about 22 0m long and the gubernaculum 18 0m. 
The cloacal opening is surrounded by irregular cuticular excrescences and anteriorly there is 
a small ventral spine. The ventral part of the precloacal body rings are more thickly 
cuticularised: they gradually reduce in thickness anterior to the level of the proximal end of 
the spicules (Fig. 9d). There is a single testis situated to the right of the gut. The female has 
two opposed, reflexed ovaries. 

DIFFERENTIAL DIAGNOSIS. Rhips paraornata sp. nov. can be distinguished from the only 
other valid species in the genus, R. ornata Cobb, 1920, by having slightly shorter R 2 cephalic 
setae, wider amphids, spicules with each part about the same length and possibly a 
differently shaped lateral differentiation. 


DISCUSSION. This is only the fifth time that valid specimens of this genus have been reported. 
Timm( 1961) described a species/?, longicauda from the Bay of Bengal, but the description is 
very poor and based only on a single immature female: it must be considered a dubious 

The original description of the type and only other species, R. ornata, was by Cobb (1920): 
the specimens from Loch Ewe are clearly similar to this species from Florida. The species 
was found again by Wieser & Hopper (1967) who provide a brief redescription and figure the 
head. Allgen (1932) found what is certainly a male Rhips species from Campbell Island and 
considered it conspecific with Cobb's species and Gerlach (1957) found R. ornata in Brazil 
but did not describe it. 

Cobb (1920) described the unusual triangular scale-like cephalic flaps but only saw the 
four sublateral ones: Wieser & Hopper (1967) correctly observed all six. The four long 
cervical setae, located at about one-third the oesophagus length in R. paraornata, were not 
reported in R. ornata but they are fine and may be lost on handling so that their absence in R. 
ornata cannot be assumed. The four sublateral cephalic flaps were positioned 'just in front of 
the ends of the amphids' in R. ornata according to Cobb (1920) and as figured by Wieser & 
Hopper (1967), whilst the ends of the amphids in R. paraornata extend further beyond the 
flaps. Both Cobb (1920) and Wieser & Hopper (1967) describe the lateral differentiation in 
the posterior portion of the body as being 'V-shaped but unfortunately provide no 
illustrations of the cuticle patterns. If the lateral differentiation is similar in both species, 
then we would not have chosen to describe the lateral differentiation as 'V'-shaped. It is 
possible, then, that the cuticle patterns are different. Finally, the spicules ofR. ornata have 
an anterior part which is 1-5 times the length of the posterior part whilst in R. paraornata the 
lengths are almost the same, measured as the arc in both cases. Therefore, although there are 
many points of similarity in overall dimensions and general anatomy, there are sufficient 
points of difference in relation to the specific sizes of cephalic setae, amphids and spicules 
and possibly the form of the cuticle lateral differentiation to substantiate the creation of a 
new species for the specimens from Loch Ewe. 

Species of the genus Rhips seem to be very closely related to Actinonema, particularly 
through the excellent redescription of the common species A. pachydermatum Cobb, 1920 
by Lorenzen (1972): both have similar amphids, cuticle patterns and the six triangular 
extensions to the cephalic shield, 'Kopfpanzer', although the flaps in A. pachydermatum are 
not as conspicuous as those in R. paraornatum and may have been overlooked by earlier 
workers. However, Actinonema does not have large double-jointed spicules and, according 
to Lorenzen (1972), between those structures which he terms spicules, but which resemble 
the lateral pieces of Rhips, there lies a single thin cuticularised tube which Lorenzen (1972) 
interprets as the cuticularised terminal portion of the vas deferens. We have studied several 
specimens of A. pachydermatum, sympatric with R. paraornata, and find the cuticularised 
tube very difficult to distinguish. However, in some specimens it is just possible to make out 
two tubes: if so, they cannot be vas deferens since A. pachydermatum is monorchic. This 
problem is of systematic importance since Rhips and Actinomena, together with a number of 
other genera including Euchromadora, are grouped together in the subfamily Euchro- 
madorinae. This subfamily was erected by Gerlach & Riemann (1973) without explanation 
but presumably because of the presence of the 'L'-shaped lateral pieces in addition to 
spicules and gubernaculum. For Actinonema to fit into this group, which seems reasonable 
through its similarity to Rhips, the spicules of Actinonema must be considered homologous 
with the lateral pieces of Rhips and either the cuticularised tube(s?) are vestigial spicules or 
spicules are absent. For the moment, this problem must remain open for further study. 


We would like to thank Dr A. D. Mclntyre of the D.A.F.S. Marine Laboratory, Aberdeen for 
providing the opportunity for one of us (Z.N.Z.) to spend two years studying benthos at that 
institute: we named one of our new species in his honour. We would also like to thank the 

246 H. M. PLATT & Z. N. ZHANG 

staff of the benthos section for help, encouragement and the provision of specimens, in 
particular Dr A. Eleflheriou and Mr D. J. Murison. 


Allgen, C. 1932. Weitere Beitrage zur Kenntnis der marinen Nematodenfauna der Campbell-insel. Nyt. 
Mag. Naturvid. 70 : 97-198. 

- 1933. Freilebende Nematoden aus dem Trondhjemsfjord. Capita zool. 4 : 1-162. 

Andrassy, I. 1976. Evolution as a basis for the systematization of nematodes. 288pp. Pitman 

London, San Francisco, Melbourne. 
Boucher, G. 1975. Nematodes des sables fins infralittoraux de la Pierre Noire (Manche occidentale). I. 

Desmodorida. Bull. Mus. natn. Hist. nat. Paris 285 : 101-128. 
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3: 1-16. 
Cobb, N. A. 1920. One hundred new nemas (type species of 100 new genera). Contrib. to a Science of 

Nematology9 : 217-343. 
De Coninck, L. A. 1942. Sur quelques especes nouvelles de Nematodes libres (Ceramonematinae Cobb, 

1933), avec quelques remarques de systematique. Bull. Mus. r. Hist. nat. Belg. 18: 1-37. 
Filipjev, I. N. 1918. Free-living marine nematodes of the Sevastopol area. Trudy osob. zool. Lab. 

Sevastop. biol. Sta. 4 : 1-350 (in Russian). 
Gerlach, S. A. 1951. Drei bemerkenswerte neue Nematoden aus der Kieler Bucht. Zool. Anz. 

147 : 37^3. 

- 1952. Nematoden aus dem Kiistengrundwasser. Abh. math, -naturw. Kl. Akad. Wiss. Mainz 

- 1956. Diagnosen neuer Nematoden aus der Kieler Bucht. Kieler Meeresforsch. 12 : 85-109. 
1957. Die Nematodenfauna des Sandstrandes an der Ku'ste von Mittelbrasilien (Brasilianische 

Meeres-Nematoden IV). Mitt. zool. Mus. berl. 33 : 41 

19630. Freilebende Meeresnematoden von den Maldiven II. Kieler Meeresforsch. 14 : 67-103. 
19636. Robbea tenax sp. n., ein merkwiirdiger mariner Nematode von den Maldiven. Int. Revue 
ges. Hydrobiol. Hydrogr. 48 : 1 53-1 58. 

& Riemann, F. 1973. The Bremerhaven checklist of aquatic nematodes. A catalogue of Nematoda 
Adenophorea excluding the Dorylaimida. Verojf.Inst. Meeresforsch. Bremerh. Suppl. 4 : 1-404. 
-, Schrage, M. & Riemann, F. 1979. Die GattungAcantholaimus (Nematoda, Chromadoridae), und 

Beobachtungen iiber einen mutmasslichen Transportmechanismus fur Spermien bie A. calathus sp. 

n. Veroff.Inst. Meeresforsch. Bremerh. 18 : 35-67. 
Haspeslagh, G. 1973. Superfamille des Ceramonematoidea (Cobb, 1933) (Nematoda), evolution et 

systematique. Annls Soc. r. zool. Belg. 102 : 235-25 1 . 
Hopper, B. E. & Cefalu, R. C. 1973. Free-living marine nematodes from Biscayne Bay, Florida V. 

Stilbonematinae: contributions to the taxonomy and morphology of the genus Eubostrichus Greef 

and related genera. Trans. Am. microsc. Soc. 94 : 578-59 1 . 
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Singer- Polignac, Paris, 19672: 29-74. 
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von Helgoland II. Desmodorida und Chromadorida. Zool. Anz. 187 : 283-302. 
1981. Entwurf eines phylogenetischen Systems der freilebenden Nematoden. Veroff. Inst. 

Meeresforsch. Bremerh. Suppl. 7 : 1-472. 
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Ass. U.K. 48: 113-142. 
& Murison, D. J. 1973. The meiofauna of a flatfish nursery ground. J. mar. biol. Ass. U.K. 

Platt, H. M. 1973. Freeliving marine nematodes from Strangford Lough, Northern Ireland. Cah. Biol. 

mar. 14:295-321. 

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135 : 239-244. 

Manuscript accepted for publication 14 September 1981 

The larval development of Crangon cmngon 
(Fabr. 1795) (Crustacea: Decapoda) 

A. R. Gurney 

Department of Zoology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 


The first larval stage of Crangon crangon (Fabr., 1795) was recorded by Du Cane in 1839 
and since then descriptions of some of the subsequent larval stages have been added. In 
1890 Ehrenbaum described five larval stages but suggested, as they were taken from 
plankton samples, that his series may not have been complete. Williamson (1901) 
reared the first three stages in the laboratory, and selected examples from tow net 
plankton samples concluding that there were probably five larval stages in all. 
Experimental evidence on the development of larvae under non-circadian light/dark 
cycles given by Dalley (1980), also suggested there were five larval stages. The present 
study is based on specimens reared in the laboratory, and describes six larval stages, one 
more than those previously recorded. 

Materials and method 

Ovigerous Crangon crangon (Fabr., 1795) were trawled in the Sound, Plymouth, in 
March, 1980. Rearing techniques follow those of Fincham, 1977. Representatives of 
each stage are deposited in the Crustacea collections of the British Museum (Natural 
History), registration numbers 1981 : 394^00. 

Description of larval stages 

Key characters are printed in italic type', many setal counts have been omitted but these 
are recorded in Table 1 . 

ZOEA 1 (Fig. 1 ) mean size 2 mm 
Head (Figs la, b): eyes are sessile. 
Carapace (Figs la, b): without spines; rostrum short. 

Antenna 1 (Fig. Ic): single peduncle segment bearing external flagellar segment with 
three aesthetascs distally plus one seta; terminal setose spine on peduncle segment. 
Antenna 2 (Fig. Id): expedite broad and flat with ten setae, all plumose except the penulti- 
mate, subapical medial spine absent; endopodite with terminal spinous seta; spine on 
distal edge of peduncle segment at base of endopodite. 
Mandibles (Fig. le): symmetrical. 

Maxilla 1 (Fig. If): coxa with seven spines; basis with five spines; endopod with five 
setae plus one small spine. 

Maxilla 2 (Figs Ig, h): endopod coxa bilobed on inner margin, proximal lobe with six setae, 
distal lobe with four setae; basis one with four setae; basis two with four setae, 
endopodite with four lobes on inner margin, fourth proximal lobe with three setae, third 
sub proximal lobe with two setae, second subdistal lobe with one seta, distal lobe with two 
setae; exopodite with five setae. 

Bull. Br. Mus. nat. Hist. (Zool.) 42 (4) : 247-262 Issued 24 June 1 982 

248 A. R. GURNEY 

Maxillipeds 1-3 (Figs li-1): with natatory exopodites. 

Pereiopod 1 (Fig. 1 m): rudimentary, biramous. 

Pereiopods 2-4 (Figs 1 m): rudimentary, uniramous. 

Pereiopod 5: absent. 

Abdomen (figs la, b): 5 somites, 6th somite continuous with telson. 

Telson (Fig. In): fans out distally, distal margin with 7+7 terminal plumose spines. 

ZOEA 2 (Fig. 2) mean size 2 -4 mm 

Head (Figs 2a, b): eyes 'stalked'. 

Antenna 1 (Fig. 2c): peduncle two segmented; internal distal margin of second segment 


Pereiopod 1 (Fig. 2m): 1- segmented endopod with one apical seta; natatory exopodite. 

Telson (Fig. 2o): distal margin with 8 + 8 terminal plumose spines. 

ZOEA 3 (Fig. 3) mean size 3'0 mm 

Antenna 1 (Fig. 3c): external jlagellum with four distal aesthetascs, one narrower than 

others and one distal seta. 

Pereiopod 1 (Fig. 3k): 2-segmented endopodite. 

Telson (Fig. 3n): divided from somite 6 by suture; exopod and endopod of uropod with 

fringing plumose setae. 

ZOEA 4 (Figs 4 & 5) mean size 3'5 mm 

Antenna 1 (Fig. 4c): rudimentary stylocerite. 

Periopod 1 (Fig. 5d): 3 -segmented endopodite. 

Abdomen (Fig. 4b): rudimentary buds of pleopods on somites 1-5. 

ZOEA 5 (Figs 6 & 7) mean size 4- 1 mm 

Antenna 1 (Fig. 6c): external jlagellum segment with 4 wide distal aesthetascs and one 

narrow sub-distal aesthetasc plus one setule. 

Pereiopod 1 (Fig. 7d): 4-segmented endopodite. 

Pereiopods 2-3 (Fig. 7e): uniramous 1 -segmented. 

Pereiopod 4 (Fig. 7f): uniramous 4-segmented. 

Pereiopod 5 (Fig. 7g): uniramous 5-segmented. 

ZOEA 6 (Figs 8 & 9) mean size 4*2 mm 
Pereiopod 1 (Fig. 9d): 5-segmented endopodite. 
Pereiopods 2-5 (Figs 9e-f): uniramous 6-segmented. 

POST LARVA 1 (Figs 10-12) mean size 4-4 mm 

Carapace (Figs lOa, b): rostrum short; one dorso-medial tooth. 

Antenna 1 (Fig. lOc): 3-segmented peduncle; external flagellum 2-segmented with four 

distal and two sub-distal aesthetascs, internal flagellum 3-segmented; stylocerite with 

terminal plumose setae. 

Antenna 2 (Fig. lOd): exopodite with 19 plumose setae; endopodite multisegmented. 

Mandibles (Fig. 10e): symmetrical. 

Maxilla 1 (Fig. 10f): coxa with 5 spines; basis with 9 spines plus one plumose seta; 

endopod with one spine. 

Maxilla 2 (Fig. lOg): endopod reduced; exopod >22 plumose setae. 

Maxilliped 1 (Fig. 11 a): endopod reduced with one plumose seta; exopod with 3 short 

plumose setae distally plus one seta, one plumose seta proximally; epipodite present. 

Maxilliped 2 (Figs, lib, c): endopod 5-segmented, exopod with reduced setae. 

Maxilliped 3 (Fig. lid): endopod 3-segmented; exopod with very reduced plumose 


Pereiopod 1 (Figs lie, 0: exopod reduced; endopod 4-segmented, propodus and dactylus 


Pereiopod 2 (Fig. 1 Ig): 5-segmented, chelate; epipodite present. 

Pereiopod 3 (Figs 1 1 h, i): 7-segmented. 



Table 1 Morphological comparison of larval stages 1-6 and post larva. (v = visible; d = developed; 
a = absent; p = present; r = rudimentary; b = biramous; re = reduced) 










2*0 mm 

2*4 mm 

3-0 mm 

3'5 mm 

4'1 mm 

4-2 mm 

4-4 mm 


No. of aesthetascs. 
















No. of segments. 









Presence of subapical medial 









Exopodite; no. of plumose setae 

incl. fine setae immediately 

after spine. 









No. of setae on coxa; 

















5 + 1 

5 + 1 

5 + 1 

5 + 1 


5 + 1 



No. of endite setae on endopodite 

lobes 1,2,3, 4; 

2 1.2.3 


basis 1 ; 








basis 2; 








coxa (prox. & distal). 

6 + 4 


7 + 4 

7 + 4 




No. of plumose setae on exopodite. 









No. of setae on endite of coxa; 
















No. of plumose setae on exopodite. 









No. of segments in endopodite. 








No. of setae on endopodite. 








Presence of natatory exopodite. 









No. of segments. 








No. of setae. 








No. of segments. 








No. of setae. 









No. of segments. 









No. of segments. 









Somite 6 distal suture present. 









No. of segments. 

















250 A.R. GURNEY 

Pereiopod 4 (Figs 1 Ij, k): 7-segmented. 

Pereiopod 5 (Figs 1 11, m): 7-segmented. 

Pleopod 1 (Fig. 12a): inner margin of exopod with 6 plumose setae, 2 apical setae, 

external margin with 4 plumose setae. 

Pleopods 2-4 (Figs 12b-d): exopod inner margin 6 plumose setae, 2 apical setae, 

external margin with 5 plumose setae. 

Pleopod 5 (Fig. 12e): exopod inner margin with 5 plumose setae, 2 apical setae, external 

margin with 4 plumose setae. 

Telson (Fig. 120: 5 + 5 spines; exopod of uropod with one spine next to subapical medial 



The larval stages of Crangon crangon (Fabr., 1795) have not previously been described 
solely from laboratory reared material. Williamson (1901) figured his larval series from 
the first three stages reared in the laboratory, supplemented by larval stages four and 
five and the post larva taken from plankton samples. All earlier records indicate five 
larval stages. In the present work six larval stages are produced consistently under 
laboratory conditions. The insertion of an additional larval stage prolonging development 
is a feature shared by other carideans (Fincham, 1977). In planktonic larvae delaying the 
onset of metamorphosis until conditions are favourable provides the maximum chance 
of survival for the mainly benthic juveniles. 

Minor differences related to the insertion of an additional larval stage become 
apparent when comparisons to Williamson's (1901) work are made. The numbers of 
aesthetascs recorded in this paper accord with Williamson up to stage four when he 
then records 6 (4 distal and 2 subdistal) compared with 5 aesthetascs (4 distal and 1 
subdistal) for stage 5; 7 (4 distal and 3 subdistal) in first post larva compared with 6 (4 
distal and 2 subdistal) in post larva recorded here. Maxilliped 1 showed variation in the 
early stages. Williamson records 10 spines on the basis in stage 1, 12 in stage 2, 13 in 
stage 3 compared with 11 in stage 1, 13 in stage 2 and 14 in stage 3 recorded here. 
Pereiopods 2-5 in the last two larval stages differed in segmentation. Williamson 
showed stage 4 pereiopod 2 as 1 -segmented, pereiopod 3 as 3-segmented and 
pereiopods 4 and 5 as 2-segmented; stage 5 pereiopods 2-5 as 7-segmented. 
Recorded in this paper stage 5 pereiopod 2 is rudimentary, pereiopod 3 is 1 -segmented, 
pereiopod 4 is 4-segmented and pereiopod 5 is 5-segmented; stage 6 pereiopods 2-5 
are 6-segmented. These differences are almost certainly related to the insertion of the 
extra larval stage after stage 2 prior to metamorphosis. 


Dalley, R. 1980. The survival and development of the shrimp Crangon crangon (L.) reared in the 
laboratory under non-circadian light-dark cycles. J. exp. mar. Biol. Ecol. 47 : 101-112. 

Du Cane, C. 1 839. Metamorphosis of Crustacea. Ann. Mag. nat. Hist. 2 : 1 78-1 8 1 . 

Ehrebaum, E. 1890. Zur naturgeschichte von Crangon vulgaris Fabr. In: Studien tiber Bau, 
Entwichklung, Lebenweise und Fanguerhdltnisse des Nordsee-Granat im Auftrage der Sektion fur 
Kusten-und Hochseefischerei des Deutschen Fischerei- Vereins. Moeser, Berlin. 124 pp. 

Fincham, A. A. 1977. Larval development of British prawns and shrimps (Crustacea: Decapods: 
Natantia). 1. Laboratory methods and a review of Palaemon (Palaeander) elegans Rathke, 1837. 
Bull. Br. Mus. nat. Hist. (Zool.)32(l) : 1-28. 

Williamson, H. C. 1901. On the larval stages of decapod Crustacea the shrimp (Crangon vulgaris, 
Fabr.) Rep. Fishery Bd Scotl. 19 : 92-1 19. 

Manuscript accepted for publication 28 September 1 98 1 


b /- \ c / I d 


Fig. 1 Zoea 1: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (f) 
maxilla 1 ; (g) maxilla 2; (h) armature of maxilla 2; (i) maxilliped 1 ; (j) armature of maxilliped 1 ; 
(k) maxilliped 2; (1) maxilliped 3; (m) pereiopods 1-5; (n) telson. 
Bar scales: a, b = 0'35 mm; c-n = 0' 1 mm. 

Fig. 2 Zoea 2: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 
maxilla 1; (g) maxilla 2; (h) armature of maxilla 2; (i) maxilliped 1; (j) armature of maxilliped 1; 
(k) maxilliped 2; (1) maxilliped 3; (m) pereiopod 1; (n) pereiopods 2-5; (o) telson; (p) detail of 
spine, 3rd from right, on telson. 
Bar scales: a, b = 0'3 mm; c-p = 0' 1 mm. 



Fig. 3 Zoea 3: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 
maxilla 1; (g) maxilla 2; (h) maxilliped 1; (i) armature of maxilliped 1; (j) maxilliped 2; (k) 
maxilliped 3; (1) pereiopod 1 ; (m) pereiopods 2-5; (n) telson. 
Bar scales: a, b = - 2 mm; c-g = 1 mm. 



Fig. 4 Zoea 4: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 
maxilla 1 ; (g) maxilla 2. 
Bar scales: a, b = - 3 mm; c-g = - 1 mm. 



Fig. 5 Zoea 4: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) 
pereiopods 2 & 3; (f) pereiopods 4 & 5; (g) telson. 
Bar scales: a-g = 0' 1 mm. 



Fig. 6 Zoea 5: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (0 
maxilla 1 ; (g) maxilla 2. 
Bar scales: a, b = 0'3 mm; c-g = - 1 mm. 



Fig. 7 Zoea 5: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) 
pereiopods 2 & 3; (f) pereiopods 4 & 5; (g) telson. 
Bar scales: a-g = 0- 1 mm. 




Fig. 8 Zoea 6: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) mandibles; (f) 
maxilla 1 ; (g) maxilla 2. 
Bar scales: a, b = 0*3 mm; c-g = 0- 1 mm. 



Fig. 9 Zoea 6: (a) maxilliped 1; (b) maxilliped 2; (c) maxilliped 3; (d) pereiopod 1; (e) 
pereiopods 2 & 3; (0 pereiopods 4 & 5; (g) telson. 
Bar scales: a-g = 0' 1 mm. 



Fig. 10 Post Larva 1: (a) dorsal view; (b) lateral view; (c) antenna 1; (d) antenna 2; (e) 
mandibles; (0 maxilla 1 ; (g) maxilla 2. 
Bar scales: a = 0*3 5 mm;b = 0-4 mm;c-g = 0-l mm. 



Fig. 11 Post Larva 1: (a) maxilliped 1; (b) maxilliped 2; (c) detail of endopod of maxilliped 2; 
(d) maxilliped 3; (e) pereiopod 1; (0 detail of pereiopod 1; (g) pereiopod 2; (h) pereiopod 3; (i) 
detail of terminal segment pereiopod 3; (j) pereiopod 4; (k) detail of terminal segment 
pereiopod 4; (1) pereiopod 5; (m) detail of terminal segment pereiopod 5. 
Bar scales: a-m = 0- 1 mm. 




be d 

Fig. 12 Post Larva 1: (a) pleopod 1; (b) pleopod 2; (c) pleopod 3; (d) pleopod 4; (e) pleopod 5; 
(0 telson. 
Bar scales: a-f = 0' 1 mm. 

A revision of the spider genus Cocalodes with a 
description of a new related genus (Araneae: 

F. R. Wanless 

Department of Zoology, British Museum (Natural History) Cromwell Road, London 
SW7 5BD. 


The genus Cocalodes Pocock, 1897, known from Amboina, Seram, Halmahera and 
New Guinea, is comprised of 12 species, six of which are described here as new. 
Petrunkevitch (1928) placed Cocalodes in the subfamily Boethinae, but as far as I am 
aware it shows no close affinities with the nominate genus Boethus or related genera 
(sensu Wanless, 1981). It is, however, closely allied to Allococalodes gen. n., proposed 
here for two new taxa from New Guinea. 

Both genera are unique in that the majority of males possess a well developed median horn 
arising from a sclerite between the lower basal margins of the chelicerae (Figs 8 A, 2 1 B). This 
structure has not been described in Cocalodes by earlier authors, for in the majority of 
species it does not protrude beyond the edge of the clypeus and is readily overlooked. The 
horn is not homologous with the paired horns ofPadilla Peckham & Peckham, which arise 
from the basal region of the anterior surface of the chelicerae. Neither is it homologous with 
the horn of Thorellia Keyserling which arises from the clypeus. 

The male palps of both Cocalodes and Allococalodes are also of interest in possessing a 
functional conductor and median apophysis which seldom occur in other Salticidae. In 
Allococalodes the functional conductor and median apophysis are lobe-like (Fig. 16E) and 
relatively simple when compared with the elaborate fan-like functional conductor and bifid 
median apophysis characteristic of Cocalodes (Fig. 6G). The less complex structures of 
Allococalodes are possibly vestigial or germinal in respect of those of Cocalodes, but for the 
present it is a matter for conjecture as the affinities of both genera are uncertain. 

The standard abbreviations and measurements are those used by Wanless (1978), 
but for the leg spination the system is that used by Platnick and Shadab (1975). 

Genus COCALODES Pocock 

Cocalodes Pocock, 1897:627. Type species Cocalodes leptopus Pocock, by original designation. 
Simon, 1901:400, 403^405. Waterhouse, 1902:80. Petrunkevitch, 1928:181. Neave, 1939, 
1 : 778. Roewer, 1954 : 936. Bonnet, 1956 : 1 172. 

DEFINITION. Small or large spiders ranging from about 4*5 to 11 -Omm in length. Most 
species elongate and narrow with long slender legs, the abdomen usually marked with 
characteristic dark lateral bands; chelicerae robust, in males usually elongate and 
porrect with a median horn which sometimes protrudes beyond the clypeus; not hirsute, 
fringes lacking. 

Carapace: longer than broad, moderately high, widest at about level of coxae II-III; 
fovea long, weakly sulciform, positioned more or less midway between posterior 
margins of posterior lateral eyes. Eyes: with black surrounds except anterior medians; 
posterior medians and posterior laterals on moderately well developed tubercles; 

Bull. Br. Mus. not. Hist. (Zool.) 42(4): 263-298 Issued 24 June 1 982 

264 F. R. WANLESS 

arranged in three transverse rows, comprised of anterior medians (AM) and anterior 
laterals (AL), posterior medians (PM), and posterior laterals (PL); anteriors more or 
less contiguous with apices procurved in frontal view and recurved in dorsal view; 
anterior medians largest; anterior laterals greater than half diameter of anterior medians; 
posterior medians relatively large, positioned closer to and on or slightly outside optical axis 
of anterior lateral eyes; posterior laterals as large or slightly smaller than anterior laterals and 
set inside lateral margins of carapace when viewed from above; quadrangle formed by 
posterior median and posterior lateral eyes broader than long and wider behind; entire 
quadrangle, measured from between bases of anterior medians to posterior margins of 
posterior laterals, occupying between 47 and 60 per cent of carapace length. Clypeus: 
between 18 and 42 per cent of diameter of anterior median eyes. Chelicerae: robust, slightly 
diverging and inclined anteriorly in females; in males usually elongate, diverging and porrect 
with median horn of variable length which arises from between the lower margins of the 
cheliceral bases (Fig. 2 1 B); pro- and retromargins usually with three teeth. Maxillae: long 
and diverging with rounded apices. Labium: oblong, generally less than half maxillae 
length. Sternum: more or less as in Figs 3B, 14G; slightly elongate scutiform. 
Coxae: I and II generally larger than HI-IV. Pedicel: short. Abdomen: long slender 
and tapering; spinnerets moderately long, posteriors longest with long apical articles, 
anteriors robust, slightly longer than more slender medians; spiracle a transverse slit 
just in front of anterior spinnerets; tracheal system not examined; position of colulus 
indicated by scant group of hairs; anal tubercle cone-shaped. Legs: long and slender; spines 
strong and numerous; claws pectinate; tufts present; scopulae absent. Female palps: long and 
slender with apical claw. Male palps: moderately complex, intra- and generally inter- 
specifically distinct. Femora long and bowed; patellae long; tibiae long with broad retro- 
lateral or ventral apophysis; cymbium with hair tuft protecting embolic tip (Fig. 2H), distal 
finger-like extension, swelling on retrolateral basal margin and usually a group of peg-like 
spines on dorsal basal margin (Figs 1 A; ISA); embolus (e) very long and slender, sometimes 
thread-like, resting distally in well developed membraneous fan-like functional conductoi r 
(c), the tip of which ends in a sclerotized spur; a bifid median apophysis arises from a trans- 
lucent pleated membrane on the tegulum (Figs 1 B; 4H); tegulum (t) irregular in form with 
seminal ducts; median hematodocha in form of a membraneous sac between tegulum and 
subtegulum only evident in fully expanded palps; subtegulum (st) a sclerotized ring at distal 
end of basal hematodocha (bh). Epigynes: of various forms; median septum sometimes 
present and occasionally bearing lateral pouches; introductory ducts long, simply looped or 
convoluted, of variable width, sometimes narrow (Fig. 8G, H), or sac-like (Figs 4F, G; 
6F) or resembling spermathecae (Fig. 141, J); spermathecae of various shapes with leaf-like 
fertilization ducts; spherical objects frequently present in interstitial spaces (Fig. 2 1C-F). 

AFFINITIES. The general habitus, presence of a median cheliceral horn, functional conductor 
and median apophysis suggest that Cocalodes and Allococalodes are closely related, even 
sister groups. Both genera are also probably allied to Holcolaetis Simon, from Africa and 
Sonoita Peckham & Peckham from South Africa, as the male papal organs evidently possess 
an homologous conductor and median apophysis. 

DIAGNOSIS. Males of Cocalodes are readily distinguished from other Oriental salticids by the 
distinctive palpal organs. Females are separated with more difficulty by the geographical 
distribution, relatively large posterior median eyes, by the structure of the epigynes and by 
having three teeth on the posterior margin of the chelicerae. A more useful diagnosis or 
assessment of affinities cannot be given until supposedly related genera have been revised. 

REMARKS. 1. The sclerite from which the median horn arises in Cocalodes and 
Allococalodes has not been routinely examined in previous revisions and not even in 
all females of Cocalodes in the present study, as the chelicerae have to be displaced or 
detatched from the carapace to expose it. A cursory examination of several Salticidae 
suggests that the sclerite is present in all members of the family. It is usually seen as 





Fig. 1 Cocalodes papuanus Simon. Expanded d 1 palp: A, retrolateral; B, prolateral. 
Abbreviations: bh, basal hematodocha; c, functional conductor; e, embolus; ma, median 
apophysis; p, peg-like cymbial spines; st, subtegulum; t, tegulum. 

266 F. R. WANLESS 

an indistinct elongate plate embedded in tissue between the bases of the lower 
margins of the chelicerae. In females of Cocalodes thoracicus Szombathy and in 
males and females of Holcolaetis it forms a peg-like horn similar to that of C. 
papuanus Simon (Fig. 2 IB). This does not, however, necessarily support a relation- 
ship between Holcolaetis and Cocalodes as our knowledge of the development of 
this sclerite is inadequate. 

2. The spherical objects found clustered or irregularly distributed in the interstitial 
spaces of the epigynes of most female Cocalodes vary in diameter from 8 to 20 //. 
Sometimes they can be seen through the cuticle of intact epigynes (Fig. 14H), but they 
are most readily observed in epigynes which have been cleared in lactic acid (Figs 
20F; 21C-F). When examined by transmitted light microscopy most spheres appear 
to have a refractile centre, but with interferance microscopy, the centres are trans- 
formed into surface depressions which resemble a lunar crater with indistinct lines 
radiating outwards from the raised margins t (Fig. 2 IF). Some spheres appear to have 
collapsed and look ragged with irregular and illdefmed craters. 

These objects, which are not known to occur in other Salticidae, resemble the 
unidentified spheres described by Forster (1980) from the epigyne of a Gamaso- 
morpha species (Family Oonopidae). A concensus of opinion (Forster, 1980) favoured a 
sporozoan infection, but this seems unlikely in the present case as the spheres lack the 
rod-like structures illustrated by Forster, furthermore their presence in almost every female 
of a wide range of species suggests they are a natural feature of these epigynes, which could be 
associated with spermatogenisis, oviposition or even plugging. Clearly, they require further 

Lists of species in the genus Cocalodes Pocock, 1897 

Cocalodes cygnatus sp. n. 

C. expers sp. n. 

C. innotabilis sp. n. 

C. leptopus Pocock, 1897 

C. longicornis sp. n. 

C. longipes (Thorell, 1881) 

C. macellus (Thorell, 1878) 

C. papuanus Simon, 1900 

C. platnicki sp. n. 

C. protervus (Thorell, 1881) 

C. turgidus sp. n. 

C. thoracicus Szombathy, 1915 

The species of Cocalodes could on the basis of the development of the epigynal 
septum be divided into two groups. However, these groups have not been proposed as 
there does not appear to be any correlation with characters of the male palpal organs. 
This may be a natural phenomenon or the result of incorrectly matching males and 
females. Matching the sexes has been difficult in the present study and where doubts 
have existed, these are mentioned in the species descriptions. As is often the case, 
additional material should resolve many of these problems. 

Key to species of Cocalodes 

Males (the males of expers, protervus and turgidus are unknown) 

1 Dorsal prong of median apophysis broad and distally truncate (Fig. 2C, F); chelicerae with 

dorsal cluster of stout bristles (Fig. 2 A) papuanus Simon (p. 267) 

- Dorsal prong of median apophysis otherwise; chelicerae lacking stout bristles 

2 Peg-like cymbial spines present (Figs 4C; ISA) 3 


Peg-like cymbial spines absent 8 

3 Conductor tip cygniform (Fig. 6G) cygnatussp. n. (p. 273) 

- Conductor tip otherwise 4 

4 Dorsal and ventral prongs of median apophysis more or less equal in length (Fig. 

IOC, D) thoracicus Szombathy (p. 280) 

- Dorsal prong of median apophysis much longer than ventral prong .... 5 

5 Dorsal prong of median apophysis finger-like or scimitar-shaped (Figs 1 3C, E; 1 5D, E) 6 
Dorsal prong of median apophysis acuminate (Figs 4C; 7H, I) 7 

6 Conductor tip forming a triangular plate (Fig. 13G); dorsal prong of median apophysis 

scimitar-shaped (Fig. 13 E) longipes (Thorell) (p. 284) 

Conductor tip otherwise (Fig. 1 5F); dorsal prong of median apophysis finger-like 

(Fig. 15D, E) innotabilis sp. n. (p. 288) 

7 Conductor tip slender with minute barb (Fig. 7G, I) .... macellus (Thorell) (p. 275) 
Conductor tip robust, barb lacking (Fig. 4H) leptopus Pocock (p. 270) 

8 Dorsal prong of median apophysis relatively slender (Fig. 14D, E); cheliceral horn 

not protruding beyond clypeal margin platnickisp. n. (p. 286) 

- Dorsal prong of median apophysis relatively robust (Fig. 8B, C); cheliceral horn 

protruding well beyond clypeal margin (Fig. 8A, D) . . . . longicornis sp. n. (p. 277) 

Females (the female of innotabilis is unknown) 

1 Epigyne with median septum 

Epigyne lacking median septum 7 

2 Epigyne with relatively large openings (Fig. 3D) papuanus Simon (p. 267) 

- Epigyne with relatively small openings 3 

3 Epigynal septum narrow and not extending to posterior margin (Fig. 7C) ... 4 

- Epigynal septum broad, extending to posterior margin 6 

4 Clypeus white haired; epigynal openings more or less ovoid in outline (Figs 9B, 7C) 5 
Clypeus not white haired; epigynal openings indistinct (Fig. 8F) . . longicornis sp. n. (p. 277) 

5 Spermathecae large and dark (Fig. 9 B) protervus (Thorell) (p. 279) 

Spermathecae small and pale, hardly if at all visible through integument (Fig. 

7C, D) macellus (Thorell) (p. 275) 

6 Epigynal septum with median bulge (Fig. 12 B) turgidus sp. n. (p. 283) 

Epigynal septum lacking median bulge (Fig. 1 1 B, C) . . . thoracicus Szombathy (p. 280) 

7 Epigyne with small median openings and broad posterior ledge (Fig. 1 4H) platnicki sp. n. (p. 286) 

- Epigyne otherwise 8 

8 Epigyne slightly depressed with comma-shaped introductory ducts (Fig. 1 3B) . 

longipes (Thorell) (p. 284) 
Epigyne otherwise 9 

9 Epigyne with deep median notch on posterior margin (Fig. 5C) . . . expers sp. n. (p. 27 1 ) 
Epigyne otherwise 10 

10 Epigyne with two notches on posterior margin (Fig. 6 D) .... cygnatussp. n. (p. 273) 
Epigyne without notches on posterior margin (Fig. 4E) . . . . leptopus Pocock (p. 270) 

Cocalodes papuanus Simon 
(Figs2A-H;3A-E; 18C,E; 19A,B;21B) 

Cocalodes papuanus Simon, 1900 : 32, rf. LECTOTYPE rf (here designated) Irian Jaya (MNHN, Paris) 

[examined]. Simon, 1901:403^04. Roewer, 1954:936. Bonnet, 1956:1173. Proszynski, 

1971 :390. 
Cocalodes armatissimus Strand, 1913:122, d. LECTOTYPE <J (here designated) Schouten 

Island (FS, Frankfurt am Main) [examined]. Strand, 1915:262. Roewer, 1954:936. Bonnet, 

1956 : 1 172. Proszynski, 1971 : 390. Syn. n. 
Cocalodes plebejus Szombathy, 1915:468, cf, 9, immatures. Syntypes (presumably in TM, 

Budapest) [not examined]. Roewer, 1954 : 936. Bonnet, 1956 : 1 173. Syn. n. 

REMARKS. Szombathy (1915) supplied good figures of the palpal organs in his original 
description of C. plebejus and there is little doubt that plebejus and papuanus are 

268 F. R. WANLESS 

DIAGNOSIS. Separated from other species of Cocalodes by the cluster of cheliceral 
setae and truncate median apophysis (Fig. 2C, F) in males, and by the large rounded 
epigynal openings in females (Fig. 3D). 

MALE from Madang, Papua New Guinea. In good condition. Carapace (Fig. 2A, B): 
pale yellow-brown grading to orange-brown in eye region with dark brown bands 
around margins and from PL's to posterior margin of thorax; pale areas clothed in 
creamy white hairs with orange-brown or black hairs elsewhere. Eyes: with black 
surrounds except AM; fringed in creamy white and pale amber hairs. Clypeus: 
clothed in very fine whitish hairs. Chelicerae: elongate, porrect and diverging; orange-brown 
with dorsal cluster of stout spines; shiny under some angles of illumination; promargin with 
four teeth, retromargin with two; cheliceral horn small, not protruding beyond clypeal 
margin. Maxillae and labium: pale yellow grading to whitish yellow along inner margins of 
maxillae. Sternum: more or less as in female; pale yellow, shiny. Coxae: pale yellow to 
whitish yellow with black lateral bands on coxae I. Abdomen: pale yellow with sooty 
markings and blackish lateral bands above, greyish black below; clothed in fine iridescent 
setae; spinnerets pale yellow heavily tinged grey-black except for whitish medians. Legs: legs 
I pale yellow to amber with blackish lateral markings on femora and metatarsi; legs II 
similar, but lighter with less conspicuous dark markings; legs III pale yellow grading to light 
amber distally; legs IV as III, but with black lateral markings on patellae, tibiae and 
metatarsi; spines numerous, strongest on legs I. Spination of legs I: metatarsi v 2-2-2, 
p 1-0-1, r 1-0-1; tibiae v 2-2-2, p 1-1-1, r 1-1-1, d 1-0-1; patellae 1-0-0, r 1-0-0; 
femora p 1-1-1, d 1-1-3, r 1-2-1. Palp (Figs 2C, E, F, H; 18C, E): the truncate dorsal 
prong of the median apophysis and pronounced functional conductor are characteristic of 
this species. 

Dimensions (mm): total length 7*88; carapace length 3*02, breadth 2-22, height 
1-44; abdomen length 4-84; eyes, anterior row 1-96, middle row 1*68, posterior row 
1-68; quadrangle length 1-6 (52% of carapace length). Ratios: AM : AL : 
PM: PL:: 16:9:6:9; AL-PM-PL :: 10-13; AM : CL (clypeus) :: 16 : 3 (18% of AM 

FEMALE (formerly undescribed) from Madang, Papua New Guinea. In good condition. 
Carapace (Fig. 3A): pale yellow-brown, shiny; clothed in fine whitish hairs with 
bands composed of amber hairs from posterior lateral eyes to posterior thoracic 
margin. Eyes: as in cf. Clypeus: below anterior median eyes and inner sides of anterior 
laterals fringed in pale amber hairs with outer sides of anterior laterals densely white 
haired. Chelicerae: robust, porrect and diverging; yellow-brown, shiny; thinly clothed in 
yellow-brown hairs; both margins with three teeth. Maxillae and labium: pale yellow, shiny. 
Sternum (Fig. 3B): pale yellow, shiny. Coxae: pale yellow, shiny. Abdomen: whitish yellow; 
irregularly clothed (?partly rubbed) in very fine iridescent setae with longitudinal bands, 
composed of dark amber hairs, flanked externally by white haired bands on the sides; 
spinnerets pale yellow tinged grey. Legs: pale yellow-brown with blackish lateral streaks on 
metatarsi IV. Spination of legs I: metatarsi v 4-0-2, p 1-0-1 , r 1-0-1 ; tibiae v 2-3-1 , p 1-1-1 , 
r 1-1-1, d 1-0-1; patellae p 1-0-0, r 1-0-0; femora p 0-2-1, d 1-1-2, r \-2-\.EpigyneQr\& 
3C-E; 19 A, B): clothed in testaceous hairs. 

Dimensions (mm): total length 10*8; carapace length 3-76, breadth 2-8, height 1-64; 
abdomen length 6*8; eyes, anterior row 2'26, middle row 1'96, posterior row 1'96; 
quadrangle length 1-8 (47% of carapace length). Ratios: AM : AL : 
PM : PL :: 19 : 10 : 5-5 : 10; AL-PM-PL :: 1 1-16; AM : CL :: 19 : 5 (26% of AM diameter). 

VARIATION, rf total length varies from 4-88 to 10*4 mm, carapace length 2-6-3-96 mm 
(seven specimens). 9 total length 8-48-10-8 mm, carapace length 2-92-3-76 mm (three 

In males the number of peg-like spines on the cymbium varies from two to four. In 
one specimen (lectotype of C. papuanus) the left palp has two spines on the cymbium while 



the right has three. In females the epigyne varies slightly, the dorsal rims of the rounded 
openings sometimes lie across the posterior margins of the spermathecae. 

DISTRIBUTION. Papua New Guinea; Irian Jaya. 

MATERIAL EXAMINED. Papua New Guinea: Madang Province, Madang, 22.iii.1979 (H. 
W. Levi, Y. Lubin, B. Robinson) (MCZ, Harvard): 19, garden, night collection; Icf, 
40 Km S. of Madang, swamp forest. Astrolabe Bay: (R. Rohde} (MNHU, Berlin. 
17795). Irian Jaya: Dorey, Lectotype rf, (A. Raffray) (MNHN, Paris. 5479); Schouten Island, 


Fig. 2 Cocalodes papuanus Simon, cf; A, dorsal; B, lateral; C, palp, retrolateral; D, 
cleared fang; E, palp, prolateral; F, median apophysis and tip of functional conductor; 
G, cheliceral teeth inner view; H, palp, ventral. 



Fig. 3 Cocalodes papuanus Simon. 9: A, dorsal; B, sternum; C, vulva ventral; D, epigyne; E, 

vulva, dorsal. 

Woges, [lectotype rf of C armatissimus], 1909, (E. Wolf) (FS, Frankfurt am Main. 2431). 
Sukarnapura (Hollandia) AMNH, New York): Irf, v. 1945 (Borys Malkin); 19, rainforest, 
250 ft. xii. 1944 (H. Hoogstraal); Irf, on foliage, rainforest, 300 ft. xii.1944 (L. W. Saylof). 
Sukarnapura: 2rf, 1936 (L. E. Cheeseman) (BMNH. 1937.12.13.117); 19, 1936 (L. E. 
Cheeseman)(]BMNH. 1937.12.13.161). 

Cocalodes leptopus Pocock 

Cocalodes leptopus Pocock, 1897:628, 9. Holotype 9, Indonesia (BMNH) [examined]. Simon, 
1901 : 403^05. Petrunkevitch, 1928: 181. Roewer, 1954:936. Bonnet, 1956: 1172. Proszynski, 
1971 :390. 

Cocalodes melanognathus Pocock, 1897:629, rf. Holotype rf, Indonesia (BMNH) [examined]. 
Simon, 1901 : 403^04. Roewer, 1954 : 936. Bonnet, 1956 : 1 172. Proszynski, 1971 : 390. Syn. n. 

REMARKS. As C. leptopus and C. melanognathus both originate from Halmahera and 
are only known from separate sexes, they are regarded here as being conspecific. 

DIAGNOSIS. From females of C. cignatus and C. expers by the apparent absence of a 
notch or notches on the posterior margin of the epigynal plate (Fig. 4E). From male 
cygnatus by the lack of a white moustache below the anterior median eyes and by the 
curved tip of the functional conductor (Fig. 4H). Males of expers are unknown. 


FEMALE HOLOTYPE. In poor condition. Carapace: brown-black with orange-brown eye 
region; irregularly clothed in whitish hairs, especially dense below lateral eyes. Eyes: 
with black surrounds; irregularly fringed in whitish hairs. Clypeus: densely white 
haired. Chelicerae: robust, inclined anteriorly and slightly diverging; brown with 
lighter brown markings; pro- and retromargins with three teeth. Maxillae (Fig. 4D): 
brownish orange grading to yellow-brown along inner margins. Labium: brownish 
orange tipped yellow-brown. Sternum: greenish yellow with thin brownish margins; 
shiny. Coxae: dark grey tinged greenish yellow. Abdomen: greenish yellow with 
brownish lateral markings and two pairs of impressed spots dorsally; clothed in 
whitish hairs (mostly rubbed on dorsum) with irregular longitudinal red haired bands 
on each side; spinnerets greenish yellow tinged black. Legs: brownish orange tinged 
greyish green, femora IV and patellae IV with black spots; spines strong and numerous. 
Spination of legs I: metatarsi v 2-0-0, p 1-1-1, d 0-0-2, r 1-1-1; tibiae v 4-4-4; patellae p 
1-0-0, r 1-0-0; femora p 1-0-2, d 0-2-2, r 0-1-1 . Epigyne (Figs 4E-G; 19C, D): a low dark 
mound with indistinct lateral openings (arrowed in Fig. 4F). 

Dimensions (mm): total length lO'O; carapace length 3'56, breadth 3'0, height 2*24; 
abdomen length 6'56; eyes, anterior row 2'52, middle row 2*12, posterior row 2*16; 
quadrangle length 1'92 (53% of carapace length). Ratios: AM : AL : 
PM:PL:: 19: 11 :7: 10; AL-PM-PL :: 13-15; AM : CL :: 19 : 9 (47% of AM diameter). 

MALE (holotype of C. melanognathus). In poor condition. Carapace (Fig. 4A, B): 
dark chocolate brown with orange-brown eye region; badly rubbed. Eyes: with black 
surrounds except AL and AM. Clypeus: irregularly and scantily clothed in fine 
whitish hairs. Chelicerae: elongate, porrect and diverging; dark brown with bluish 
sheen under some angles of illumination; pro- and retromargins with three teeth; 
horn small, dislocated by a pin pushed through the specimen, a method frequently 
used by Pocock for orientating larger spiders. Maxillae, labium, sternum and coxae: 
more or less as in 9. Abdomen: similar to 9. Legs: broken and detached, otherwise 
similar to 9. Palp (Fig. 4C, H): similar to that of C. cygnatus, but readily distinguished 
by the curved tip of the functional conductor. 

Dimensions (mm): total length 8'8; carapace length 3*4, breadth 2'72, height 2'08; 
abdomen length 5'36; eyes, anterior row 2*48, middle row 2*08, posterior row 2*16; 
quadrangle length l - 94 (57% of carapace length). Ratios: AM : AL : 
PM:PL:: 19-5: 11-5 ::7: 11; AL-PM-PL:: 12-15; AM : CL ::19'5 : 5 (28% of AM 

DISTRIBUTION. Indonesia: Halmahera. 

MATERIAL EXAMINED. Halmahera: Patani, holotype 9, ii.1894 (Kukenthal); Soah Konorah, 
holotype rf [of C. melanognathus] 1 894 (Kukenthal) (BMNH. 1981.1.22.1-2.) 

Cocalodes expers sp. n. 
(Fig. 5A-C) 

DIAGNOSIS. From C. leptopus and C. cygnatus by the presence of a deep median notch 
on the posterior margin of the epigynal plate (Fig. 5C). 

MALE. Unknown. 

FEMALE HOLOTYPE. In good condition. Carapace (Fig. 5A, B): orange-brown with 
vague darker markings; irregularly clothed in short, fine shinning hairs which appear 
whitish or pale amber under varying angles of illumination. Eyes: with black surrounds 
except AM; fringed in whitish and pale amber hairs. Clypeus: densely clothed in creamy 
white hairs. Chelicerae: robust, divergent and inclined anteriorly; orange-brown, thinly 
clothed in clear amber hairs; pro- and retromargins with three teeth. Maxillae and labium: 
amber, shiny. Sternum: light amber with darker margins; thinly covered in light brownish 




Fig. 4 Cocalodes leptopus Pocock. 3 (holotype of C. melanognathus): A, dorsal; B, 
lateral; C, palp, retrolateral; H, palp, ventral. Holotype 9: D, sternum, coxae and mouth 
parts; E, epigyne; F, vulva, ventral; G, vulva, dorsal. 

hairs. Coxae: light amber. Abdomen: pale orange-brown with fine shinning hairs, scattered 
spots composed of amber hairs, and dark reddish brown lateral bands covered in dark amber 
hairs; spinnerets pale orange-brown. Legs: orange-brown with blackish apices on femora IV; 
spines numerous. Spination of legs I: metatarsi v 2-2-2, p 1-0-1, r 1-0-1; tibiae v 2-2-2, 
p 1-1-1, d 0-1-0, r 1-1-1; patellae p 0-1-0, r 0-1-0; femora p 2-2-1, d 1-1-3, r 2-1-1. 
Epigyne (Fig. 5C): similar to that of C. leptopus, but readily separated by the median notch 
on the posterior margin of the epigynal plate. 




A /7BT C 

Fig. 5 Cocalodes expers sp. n. Holotype 9: A, dorsal; B, carapace lateral; C, epigyne. 

Dimensions (mm): total length 10'9; carapace length 4-5, breadth 3*8, height 2*8; 
abdomen length 6*24; Eyes, anterior row 3*04, middle row 2-68, posterior row 2*68; 
quadrangle length 2*28 (50% of carapace length). Ratios: AM : AL : 
PM : PL :: 22 : 14 : 8 :14; AL-PM-PL :: 14 : 19; AM : CL :: 22 : 8 (36% of AM diameter). 

DISTRIBUTION. Papua New Guinea. 

MATERIAL EXAMINED. Papua New Guinea, D'Entrecasteaux Is., Fergusson Island, lamelele 
about 1*5 miles from Seymour Bay, 15m., camp 3. holotype 9, 1956 (Fifth Archbold Exp. to 
New Guinea, L. J. Brass) (AMNH, New York). 

REMARKS. This species has been recorded from the same locality (Fergusson Island) 
as C. longicornis sp. n. 

Cocalodes cy gnat us sp. n. 

DIAGNOSIS. From females of C. leptopus and C. expers by the presence of two small 
notches on the posterior margin of the epigynal plate (Fig. 6D). From male leptopus 
by the white moustache below the anterior median eyes and by the cygniform tip of 
the functional conductor (arrowed, Fig. 6G). Males of expers are unknown. 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 6A, B): orange-brown with dark 



Fig. 6 Cocalodes cygnatus sp. n. Holotype rf: A, dorsal; B, carapace, lateral; C, palp, 
retrolateral; G, palp, ventral. Paratype 9: D, epigyne; E, vulva, ventral; F, vulva, dorsal. 

brown margins and vague bands behind PL's; thinly clothed in whitish and pale 
amber hairs with scanty white haired patches on posterior declivity. Eyes: with black 
surrounds except AM; fringed by white hairs. Clypeus: densely white haired below 
AM. Chelicerae: elongate, porrect and diverging; dark reddish brown; shiny; thinly 
covered in pale amber hairs; pro- and retromargins with three teeth; horn small, not 
protruding beyond clypeal margin. Maxillae: brownish with inner margins orange- 
brown. Labium: dark brown tipped orange-brown. Sternum: yellow-brown with darker 
margins. Coxae: yellow-brown with black lateral stripes on coxae I. Abdomen: mottled 
grey-black with creamy white dorsal markings; spinnerets yellow-brown tinged black. Legs: 
legs I femora brown-black with yellow-brown markings, patellae and tarsi yellow-brown, 
tibiae dark brown with yellow-brown annuli, metatarsi basally yellow-brown grading to dark 
brown distally; other legs similar, but markings becoming paler except for brown-black 
blotches on femora IV; spines strong and numerous. Spination of legs I: metatarsi v 2-0-0, 
p 1-1-1, d 0-1-2, r 1-1-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0, r 1-1-1; patellae p 1-0-0, 


r 1-0-0; femora p 1-1-2, d 0-2-2, r 0-1-1. Palp (Fig. 6C, G): the tip of the functional 
conductor is sometimes obscured by the tip of the median apophysis. Also, in lateral view 
the ventral prong of the median apophysis is hardly evident, c.f. (C. leptopus). 

Dimensions (mm): total length c. 8 - 2; carapace length 3' 16, breadth 2 - 6, height 2-0; 
abdomen length 5'0; eyes, anterior row 2-36, middle row 1'98, posterior row 2-04; 
quadrangle length 1'84 (58% of carapace length). Ratios: AM:AL: 
PM:PL:: 19: 11 : 5-5 : 10; AL-PM-PL :: 1 1-15; AM : CL :: 19 : 8 (42% of AM diameter). 

FEMALE PARATYPE. In poor condition. Carapace: generally yellow-brown with blackish 
bands from PL's to thoracic margin; sparsely and irregularly clothed in creamy white hairs. 
Eyes: with black surrounds except AM; fringed in silky white hairs. Chelicerae: robust, 
diverging and inclined anteriorly; amber; shiny under some angles of illumination; with 
scattered pale amber hairs along inner margins; pro- and retromargins with three teeth. 
Maxillae, labium, sternum and coxae: yellow-brown. Abdomen: pale yellow-brown; clothed 
in fine whitish hairs with irregular lateral bands composed of amber hairs; spinnerets pale 
yellow-brown. Legs: generally yellow-brown, but with black spots on patellae IV and femora 
IV; spines strong and numerous on anterior legs becoming weaker and fewer on posteriors. 
Spination of legs I: metatarsi v 2-0-0, p 1-1-1, d 0-1-0, r 1-1-1; tibiae v 2-2-2, p 1-1-1, 
d 1-0-1, r 1-1-1; patellae r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-1-1. Epigyne (Figs 
6D-F;19E, F): a low mound as is leptopus, but pale and with more apparent detail. 

Dimensions (mm): total length c. 8'0; carapace length 3' 16, breadth 2-68, height 
2*0; abdomen length 4-8; eyes, anterior row 2*44, middle row 2'07, posterior row 
2-10; quadrangle length 1-88 (59% of AM diameter). Ratios: AM:AL: 
PM: PL:: 19: 11-5:6: 11; AL-PM-PL :: 12 : 15; AM:CL::19:c. 8 (c. 42% of AM 

VARIATION. Paratype cf measures c. 7-2 mm total length, 2-8 mm carapace length. 
DISTRIBUTION. Indonesia: Halmahera Island. 

MATERIAL EXAMINED. Halmahera Island: Edkor, holotype rf, paratype $ (MNHN, 
Paris. 7682). Paratype cf, same data as holotype (BMNH, 198 1 .5. 14.1). 

REMARKS. Simon misidentified the above specimens as C. leptopus, but as far as I am 
aware there has been no reference to them in the literature. 

Cocalodes macellus (Thorell) 
(Figs 7A-I; 20A, B) 

Cocalus macellus Thorell, 1878:287, 311, 9 and juvenile. Holotype $, juvenile, Amboina 
(MCSN, Geneva) [examined]. Thorell, 1881 : 493, 706. Simon, 1901 : 407. Roewer, 1954:934. 
Bonnet, 1956: 1 173. Proszyriski, 1971 : 391. Wanless, 1981 : 256. 

DIAGNOSIS. From other species of Cocalodes by the long slender prongs of the median 
apophysis (Fig. 7H, I) in males, and the appearance of the epigyne (Figs 7C-F; 
20A, B) in females. 

HOLOTYPE FEMALE. In fair condition. Carapace: yellow-brown with pale yellowish 
eye region. Eyes: with black surrounds except AM; fringed in whitish hairs. Clypeus: 
densely white haired. Chelicerae: yellow-brown thinly clothed in testaceous hairs; 
pro- and retromargins with three teeth. Maxillae: yellow-brown with pale yellow 
blades. Labium: yellow-brown tipped yellow. Sternum: yellow-brown. Abdomen: long and 
tapered; pale yellow. Legs: generally yellow-brown; spines strong and numerous on legs I 
becoming weaker and fewer on posterior legs. Spination of legs I: metatarsi v 2-2-2, p 1-0-0, 
d 0-2-2, r 1-0-0; tibiae v 2-4-0, p 1-1-2, d 1-1-0, r 1-1-2; patellae p 1-0-0, r 1-0-0; femora 
p 1-0-1 , d 1-1-3, r 0-1-0. Epigyne (Fig. 7D): somewhat translucent; the ducts will probably 
be less evident in freshly preserved specimens. 



Dimensions (mm): total length c. 7'8; carapace length 2*88, breadth 2'34, height 
1'72; abdomen length 4'88; eyes, anterior row 2'2, middle row l - 92, posterior row 
2-0; quadrangle length 1*64 (56% of carapace length). Ratios: AM : AL : 
PM : PL :: 17 : 1 1 : 6 : 10; AL-PM-PL :: 9-12; AM : CL :: 17 : 3-5 (20% of AM diameter). 

MALE (formerly undescribed) from Draeger Harbor, Papua New Guinea. In fair 
condition. Carapace (Fig. 7A, B): dark mahogany grading to orange-brown in eye 
region; irregularly clothed in short amber hairs (?rubbed) with white haired marginal 
band from level of coxae II to IV. Eyes: with black surrounds except AM; fringed by 
shining pale amber, and white hairs. Clypeus: fringed in light brownish hairs. Chelicerae: 
moderately elongate and porrect, more or less parallel; dark orange-brown, lighter distally; 
thinly clothed in fine light brownish hairs; teeth not examined; horn not evident. Maxillae: 
dark orange-brown with yellow-brown blades. Labium: dark orange-brown. Sternum: 
yellow-brown with amber margins, shiny; sparsely clothed in fine clear hairs. Coxae: coxae I 
yellow-brown below, dark mahogany above; other coxae yellow-brown. Abdomen: badly 



Fig. 7 Cocalodes macellus (Thorell). d: A, dorsal; B, carapace, lateral; G, palp, ventral; 
H, median apophysis; I, palp, retrolateral. Holotype 9: D, epigyne. C, epigyne of 
another specimen; E, vulva, ventral; F, vulva, dorsal. 


rubbed; yellow-brown with a dark patch anteriorly and ventral grey longitudinal band. Legs: 
legs I with tarsi and patellae yellow-brown, metatarsi yellow-brown suffused with black 
distally, tibiae dark mahogany with apices yellow-brown, femora dark mahogany; legs II-III 
yellow-brown; legs IV missing; spines strong and numerous. Spination of legs I: 
metatarsi v 2-2-2, p 1-0-0, d 1-2-2, r 1-0-0; tibiae v 3-2-4, p 0-1-1, d 1-1-0, 
r 1-1-1; patellae p 1-0-0, r 1^0-0; femora p 0-1-1, d 0-2-2. Palp (Fig. 7G-I): femora 
brown-black with distal third light yellow, other segments light yellow; both prongs of 
median apophysis relatively long and slender. 

Dimensions (mm): total length 5'68; carapace length 2*32, breadth 1'92, height 
1-48; abdomen length 3'2; eyes, anterior row 1-85, middle row 1'56, posterior row 
1-72; quadrangle length 1-4 (60% of carapace length). Ratios: AM : AL : 
PM : PL :: 15 : 9 : 4 : 8-5; AL-PM-PL :: 9*5-10; AM : CL :: 15 : 4-5 (30% of AM diameter). 

VARIATION. 9 from Papua New Guinea measures c. 6*9 mm total length, 2-76 mm 
carapace length. 

The epigyne (Fig. 7C) is slightly less translucent and the looped ducts are less 

DISTRIBUTION. Indonesia: Amboina; Papua New Guinea. 

MATERIAL EXAMINED. Amboina: holotype 9, juvenile, (O. Beccari) (MCSN, Geneva). Papua 
New Guinea: Draeger Harbour, 5 miles E. of Finschhafen, 19, vi.1944 (G. H. Penn) (AMNH, 
New York); Nr. River Song, 6 miles W. of Finschhafen, Icf, viii-ix. 1944, (T. F. Delaney) 
(AMNH, New York). 

Coca I odes longicornis sp. n. 

DIAGNOSIS. From males of C. longipes and C. innotabilis by the absence of peg-like 
cymbial spines, and from male C. platnicki by the thickened dorsal prong of the 
median apophysis (Fig. 8B, C). Females, presumed to be of this species, most closely 
resemble females of C. macellus, but may be separated by the lack of a dense covering 
of white hairs on the clypeus and by the laterally extended introductory ducts (Figs 
8F, G, H; 2 1 A) of the epigynum. 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 8A, D): dark rufose with eye 
region dark amber; sparsely clothed in short fine amber hairs with whitish hairs in 
foveal region and a narrow marginal band of white hairs extending from level of coxae 
II to IV. Eyes: with black surrounds except AM; fringed by shining amber hairs with 
white hairs outside AL's. Clypeus: densely white haired below AM, sparsely amber 
haired below AL. Chelicerae: elongate, porrect and diverging; rufose; shiny; pro- and 
retro-margins evidently with three teeth; horn well developed and protruding well 
beyond clypeal margin. Maxillae: orange-brown grading to yellow-brown along inner 
margins. Labium: orange-brown. Sternum: orange-brown with darker margins; sparsely 
clothed in coarse black hairs marginally with fine pale brown hairs centrally. Coxae: 
orange-brown. Abdomen: dull orange-brown with blackish lateral bands above and central 
blackish band below; spinnerets orange-brown. Legs: legs I-II dark rufose with orange- 
brown tarsi, other legs rufose grading to orange-brown distally; spines strong and numerous. 
Spination of legs I: metatarsi v 2-2-2, p 1-0-1, d 0-1-0, r 1-0-1; tibiae v 2-2-2, p 1-1-1, 
d 1-1-0, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora r 1-1-1, d 0-2-3, p 1-1-1. Palp (Fig. 
8B, C, E). 

Dimensions (mm): total length 10' 16; carapace length 4*28, breadth 3'5, height 
2-64; abdomen length 5'84; eyes, anterior row 2*84, middle row 2*48, posterior row 
2-5; quadrangle length 2-12 (49% of carapace length). Ratios: AM : AL : 
PM :PL:: 21 : 13-5:8: 13; AL-PM-PL :: 12-18; AM : CL :: 21 : 7-5 (35% of AM 



hEMALE PARATYPE. In fair condition. Carapace: orange-brown with reddish brown 
eye region and dark red-brown markings from PL's to thoracic margin; clothed in fine 
shining hairs (mostly rubbed). Eyes: with black surrounds except AM; fringed by 
whitish and pale amber hairs. Clypeus: fringed in whitish hairs. Chelicerae: robust, 
inclined anteriorly and slightly diverging; reddish with darker markings; shiny; sparsely 
fringed in pale amber hairs; pro- and retromargins with three teeth. Maxillae and labium: 
amber. Sternum: amber with darker margins; shiny; thinly clothed in fine light amber hairs. 
Abdomen: long and tapered; dull yellow-brown with blackish lateral markings; spinnerets 
yellow-brown. Legs: legs I amber with darker metatarsi; other legs amber to pale amber with 
black spots on femora IV; spines strong and numerous. Spination of legs I: metatarsi v 
2-2-2, p 1-0-1 , d 0-1-1 , r 1-0-1 ; tibiae v 3-2-2, p 0-1-1 , d 0-1-0, r 1-1-1 ; patellae p 1-0-0, 

Fig. 8 Cocalodes longicornis sp. n. Holotype d: A, dorsal; B, palp, retrolateral; C, 
median apophysis; D, carapace and chelicera, lateral; E, palp, ventral. Paratype 9: F, epigyne; 
G, vulva, ventral; H, vulva dorsal. 



r i-O-O; femora p 1-1-1, d 0-2-3, r 0-1-0. Epigyne (Fig. 8F-H; 21 A): relatively simple, the 
openings lead into narrow introductory ducts which extend and loop laterally before joining 
the spermathecae. 

Dimensions (mm): total length c. 6-4; carapace length 3'32, breadth 2'88, height 
2-08; abdomen length c. 3'0; eyes, anterior row 2'6, middle row 2-32, posterior row 
2-44; quadrangle length 2*0 (60% of carapace length). Ratios: AM : AL : 
PM : PL :: 19 : 12 : 7 : 12; AL-PM-PL :: 13 : 15-5; AM : CL :: 19 : 5 (26% of AM diameter). 

VARIATION, <S total length varies from lO'O to 10-16 mm, carapace length 3-76-4'16mm 
(three specimens). Another 9 measures c. 8'0 mm total length, 3*0 mm carapace length. 

DISTRIBUTION. Papua New Guinea. 

MATERIAL EXAMINED. Papua New Guinea: D'Entrecasteaux Islands, Normanby Island, Mt. 
Pabinama, about 3 miles ENE of Cape Prevost, 820m, camp 2, 1956 (J. L. Brass, 5th 
Archbold Exp., to New Guinea) holotype d 1 , (AMNH, New York), paratype d (BMNH. 
1981.2.2.1); Fergusson Island, Agamoia about 3 miles S. of Lake Ruaba, 200m, camp 5, 
18-24. vi.1956 (J. L. Brass, 5th Archbold Exp., to New Guinea) paratype d 1 , paratype 9, 
(AMNH, New York); New Britain, Ralum, (nr. Kokopo) Lowan, on tree trunk, paratype 9, 
27.xii. 1 896 (F. Dahl) (MNHU, Berlin. 1 7799). 

Cocalodes protervus (Thorell) 

Cocalus protervus Thorell, 1881 :493, 9. Holotype 9, West Irian (MCSN, Geneva) [examined]. 
Simon, 1901:407. Roewer, 1954:935. Bonnet, 1956:1174. Proszynski, 1971:391. Wanless 
1981 :256. 


Fig. 9 Cocalodes protervus (Thorell). Holotype 9: A, dorsal; B, epigyne. 

280 F. R. WANLESS 

DIAGNOSIS. Similar to C. macellus and C. longicornis from which it differs by the 
large darkened spermathecae (Fig. 9B). 

MALE. Unknown. 

FEMALE HOLOTYPE. In poor condition. Carapace (Fig. 9A): brownish with scattered 
white hairs, rubbed. Eyes: with black surrounds except AM; fringed by whitish hairs. 
Clypeus: densely white haired. Chelicerae: robust, inclined anteriorly and slightly 
diverging; brownish with sooty markings; sparsely clothed in light brown hairs; pro- 
and retromargins with three teeth. Maxillae and labium: pale brown. Sternum: pale 
brown with darker margins. Coxae: generally brownish. Abdomen: greyish yellow 
with darker somewhat mottled longitudinal lateral markings; clothed in short recumbent 
amber hairs. Legs: dark brown; spines long, slender and numerous. Spination of legs I: 
metatarsi v 2-2-2, p 1-2-1, d 1-0-0, r 1-0-1; tibiae v 4-3-3, p 0-2-1, d 1-1-0, r 0-0-1; 
patellae p 1-0-0, r 1-0-0; femora p 0-1-1 , d 0-2-3, r 0-2-0. Epigyne (Fig. 9B): rather dark, 
openings separated by distinct septum with black disc-like spermathecae posteriorly. 

Dimensions (mm): total length 6*8; carapace length 3*04, breadth 2-46, height 1-84; 
abdomen length 3'84; eyes, anterior row 2-32, middle row 1*98, posterior row 2*08; 
quadrangle length 1-72 (56% of carapace length). Ratios: AM:AL: 
PM: PL ::18: 11:6: 10; AL-PM-PL:: 11-13; AM : CL :: 18 : c. 7 (c. 38% of AM 


MATERIAL EXAMINED. West Irian: Pulo Faor, holotype d, 1872, (L. M. D'Albertis) 
(MCSN, Geneva). 

Cocalodes thoracicus Szombathy 
(Figs 10A-E; 1 1 A-E; 18D; 20C, D) 

Cocalodes thoracicus Szombathy, 1915:470, cf, 9. Syntypes, Papua New Guinea (?in 
Termeszettudomanyi Muzeum, Budapest) [not examined]. Roewer, 1954 : 936. Bonnet, 1956 : 1 173. 

REMARKS. Although it was not possible to examine the type specimens of C. thoracicus it 
may be recognized with a fair degree of confidence from Szombathy 's original figures, which 
show the distal cheliceral lobes (arrowed, Fig. 10A) to be more pronounced in this species 
than in others of the genus. 

DIAGNOSIS. From other species of Cocalodes by the more or less equal development of 
the prongs of the median apophysis (Figs IOC, D; 18D) in males, and by the relatively 
broad epigynal septum (Fig. 1 1 B, C) in females. 

MALE from Morobe Province. In good condition, but legs detached. Carapace brown- 
black with dark orange-brown eye area and central yellow-brown stripe from foveal 
region to posterior margin; clothed dorsally in fine shining pale amber and whitish 
hairs with short coarse black hairs around thoracic margins. Eyes: with black surrounds 
except AM; fringed by pale amber and whitish hairs. Clypeus: sparsely covered in pale 
amber and blackish hairs. Chelicerae: elongate and porrect with distal lobes; black with 
bluish metallic sheen under some angles of illumination; proximally sparsely clothed in stiff 
black hairs; promargin with four teeth, retromargin with two; horn protruding just beyond 
clypeal margin. Maxillae: brownish black grading to yellow-brown distally. Labium: 
brown-black tipped yellow-brown. Sternum: yellow-brown faintly tinged black with 
brownish orange margins. Coxae: pale yellow. Abdomen: with four impressed spots; mottled 
grey-black with a dorsal greyish yellow band having vague sooty markings; ventrally a 
longitudinal black stripe; irregularly clothed in minute setae with scattered long blackish 
hairs; spinnerets mottled yellow-brown and black. Legs: legs I tarsi pale yellow, metatarsi 
black with pale yellow basal region, tibiae black distally grading to dark yellow- 
brown to black proximally, patellae black distally grading to yellow-brown proximally, 



Fig. 10 Cocalodes thoracicus Szombathy. d: A, dorsal; B, carapace, lateral; C, palp, 
retrolateral; D, median apophysis; E, palp, ventral. 

femora yellow to yellow-brown; legs II as I, but tibiae and patellae yellow-brown; legs III 
yellow-brown to pale yellow-brown; legs IV similar, but with vague darker markings; spines 
strong and numerous. Spination of legs I: matatarsi v 2-2-2, p 1-0-0, r 1-0-1 ; tibiae v 2-2-2, 
p 1-1-1, d 1-1-0, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 1-1-2, d 0-2-1, r 0-1-2. Palp 

Dimensions (mm): total length 9*28; carapace length 3-48, breadth 3*16, height 
2'28; abdomen length 5'36; eyes, anterior row 2*56, middle row 2*28, posterior row 
2-37; quadrangle length 2*08 (59% of carapace length). Ratios: AM : AL : 
PM : PL :: 20 : 12 ; 8 : 12; AL-PM-PL :: 13-15-5; AM : CL :: 20 : 5 (25% of AM diameter). 

FEMALE from Madang Province. In good condition. Carapace (Fig. 11 A): yellow- 
brown with orange-brown eye region, a thin dark brown border line, lateral brownish 
stripes and dark brown bands from PL's to thoracic margin; clothed in very fine 
shining amber and whitish hairs. Eyes: with black surrounds except AM; fringed in 
shining pale amber and whitish hairs. Clypeus: sparsely covered in white hairs. 
Chelicerae: robust, inclined anteriorly and diverging; yellow-brown; shiny; thinly clothed in 
testaceous hairs; promargin with four teeth, retromargin with three. Maxillae and labium: 
pale yellow-brown. Sternum: pale yellow-brown with darker margins; sparsely clothed in 

282 F. R. WANLESS 

testaceous hairs. Coxae: pale yellow-brown. Abdomen: similar to d" except for pale yellow 
venter and covering of dark amber hairs over lateral bands. Legs: legs I yellow-brown with 
darker markings and a black streak on inside of femora; legs II yellow-brown with brownish 
markings; legs III yellow-brown; legs IV yellow-brown with brownish markings and black 
spots on femora and patellae; spines strong and numerous. Spination of legs I: metatarsi v 
2-2-2, p 1-0-1, r 1-0-1; tibiae v 2-2-2, p 1-1-2, r 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 
0-2-2, d 0-2-1, r 0-2-0. Epigyne (Figs 1 1B-E; 20C, D): relatively simple, but variable; the 
obscure openings (arrowed, Fig. 1 1 D) lead into wide ducts which narrow and loop 
posteriorly before entering the spermathecae. 

Dimensions (mm): total length 9'28; carapace length 3*32, breadth 2'84, height 
1-96; abdomen length 5-6; eyes, anterior row 2-44, middle row 2'2, posterior row 
2-28; quadrangle length 1*92 (57% of carapace length). Ratios: 
AM : AL: PM : PL :: 18: 12:7: 1 1; AL-PM-PL :: 12-5-15; AM : CL :: 18 : 5 (27 per cent of 
AM diameter). 

VARIATION, d 1 total length varies from 6*9 to 11 -5 mm, carapace length 2-72-4-4 mm 
(14 specimens). 9 total length 7-28-10-8 mm, carapace length 2-84^-08 mm (nine 

The cheliceral horn, possibly an allometric growth character, only protrudes well 
beyond the clypeus in large specimens. The epigynal septum varies from rectangular 
to triangular in outline (Fig. 1 1 B, C) and the rod-like extensions of the spermathecae 
(in reality part of the introductory ducts), are not always evident in uncleared 

DISTRIBUTION. Papua New Guinea; Irian Jaya. 

W. :-^c- 'i' ' JtfaMttb ..-..' - - ~~Zi&r / 

Fig. 11 Cocalodes thoracicus Szombathy. 9: A, dorsal; B, epigyne: C, epigyne of another 
specimen; D, vulva, ventral; E, vulva, dorsal. 



MATERIAL EXAMINED. Papua New Guinea: Morobe Province, Wau: Ip, iv.1979 (H. W. 
Levi); 19, iv.1979 (M. Robinson); 19, iv.1979, beating vegetation in forest, (H. W. 
Levi); 3dtf, 19, Me Adam Park, 4.iv.l979 (H. W. Levi, Y. Lubin, M. Robinson); 
Madang Province, 25 Km, N. of Madang, teak forest, night collection, Id 1 , 21.iii.1979 
(H. W. Levi, Y. Lubin. B. Robinson) (MCZ. Harvard); Draeger Harbour: 5 miles E. of 
Finschhafen, Id 1 , vi.1944 (G. H. Penn) (AMNH, New York); Maffm Bay: Id 1 , xii.1944 
(Lt. Geo. B. Sirotiak) (AMNH. New York); Astrolabe Bay, 19, (R. Rohde) (MNHN, 
Berlin. 17794); Ramu, Id 1 , (F. Dahl, Ramu Expedition) (MNHN, Berlin. 17801); Jagei 
River, Id, 2.viii.l896 (Lauterbach) (MNHN, Berlin, 17796). Irian Jaya: Sukarnapura 
(Hollandia): 2dtf, 5. i. 1945 (L. W. Saylor); Icf, iv.1945 (Borys Malkin); Id, 14.V.1945 
Borys Malkin); rain forest, 250ft. 2dtf, xii.1944 (H. Hoogstraat) (AMNH, New York); 
Humboldts Bay, 200ft. 19, iv.1936 (L. E. Cheesman) (BMNH. 1937.12.13.425); Cyclops 
Mts.Sabron,2,200ft. I $ ,v.l936 (L. E. Cheesman) (BMNH. 1937.12.13.541). 

Coca lodes turgidus sp. n. 


DIAGNOSIS. Like C. thoracicus, but differs by the median swelling of the epigynal 
septum (Fig. 12B). 

MALE. Unknown. 

FEMALE HOLOTYPE. In fair condition. Carapace (Fig. 12 A): yellow-brown with orange-brown 
eye region, dark brown lateral margins and dark brown stripes below the lateral eyes to 
posterior margin; clothed in minute creamy white setae with light brownish hairs around 
thoracic margins. Eyes: with black surrounds except AM; irregularly fringed by creamy 


Fig. 12 Cocalodes turgidus sp. n. Holotype 9: A, dorsal; B, epigyne. Paratype 9: C, vulva, 

ventral; D, vulva, dorsal. 

284 F. R. WANLESS 

white hairs. Chelicerae: robust, inclined anteriorly and diverging; amber with brownish 
markings; shiny; thinly clothed in clear pale amber hairs; pro- and retromargins with three 
teeth. Maxillae: yellow-brown with inner distal margins whitish yellow. Labium: 
yellow-brown. Sternum: light yellow-brown with darker margins; shiny; thinly clothed in 
testaceous hairs. Coxae: pale yellow-brown. Abdomen: with four impressed spots; pale 
yellow with black lateral stripes and a black ventral band. Legs: legs I with tarsi 
yellow-brown, metatarsi blackish grading to light orange-brown proximally, tibiae 
yellow-brown with blackish annuli, patellae yellow-brown, femora yellow-brown with 
blackish streaks; other legs similar, but darker markings less extensive especially on legs 
III-IV; spines strong and numerous. Spination of legs I: metatarsi v 2-2-2, p 1-0-0, d 0-1-0, 
r 1-0-1 ; tibiae v 2-2-2, p 0-1-1 , d 0-1-0, r 1-1-1 ; patellae p 1-0-0, r 1-0-0; femora p 1-1-0, 
d 0-2-3, r 0-1-1. Epigyne (Figs 12B-D;20E, F). 

Dimensions (mm): total length 8*72; carapace length 3'32, breadth 2'88, height 
1'96; abdomen length 5'28; eyes, anterior row 2*44, middle row 2- 18, posterior row 
2-32; quadrangle length 2*02 (60% of carapace length). Ratios: AM : AL : 
PM : PL :: 19 : 12 : 8 : 12; AL-PM-PL :: 1 1-16; AM : CL :: 19 : 5 (26% of AM diameter). 

VARIATION. A paratype 9 measures 8*64 mm total length, 3'28 mm carapace length. 

MATERIAL EXAMINED. Irian Jaya: Sansapor, holotype 9, paratype 9, viii-ix.1944 (R. B. 
Burrows, A.P.O. 1 59) (AMNH, New York). 

Cocalodes longipes (Thorell) 
(Fig. 13A-G) 

Cocalus longipes Thorell, 1881:494, 707, 9, d. LECTOTYPE 9, Yule Isl. PARALECTO- 
TYPE cf, Ceram Isl. (here designated) (MCSN, Geneva) [examined]. Simon, 1901:407. 
Roewer, 1954 : 935. Bonnet, 1956: 1 173. Proszyriski, 1971 : 391. Wanless 1981 : 256 

DIAGNOSIS. From other species of Cocalodes by the scimitar-shaped dorsal prong of the 
median apophysis (Fig. 13E) in males, and by the comma-like introductory ducts (Fig. 13B) 
in females. 

FEMALE LECTOTYPE. In fair condition. Carapace (Fig. 13D, F): orange-brown with 
sooty markings; irregularly clothed in short recumbent white hairs. Eyes: with black 
surrounds except AM; fringed by whitish hairs. Clypeus: thinly clothed in fine 
whitish hairs. Chelicerae: robust, inclined anteriorly and slightly diverging; dark amber; 
thinly clothed in white and light amber hairs; pro- and retromargins with three teeth. 
Maxillae: orange-brown with lighter inner margins. Labium: orange-brown. Sternum: amber 
with darker margins. Coxae: generally yellowish orange to amber. Abdomen: pale 
yellow-orange with faint sooty lateral markings outlining a central pale band; spinnerets pale 
yellow-brown. Legs: brownish amber grading to light amber distally; spines numerous and 
robust. Spination of legs I: metatarsi v 2-1-1, p 1-1-1, d 0-0-2, r 1-0-0; tibiae v 4-4-2, 
p 0-1-1, d 0-1-0; patellae r 1-0-0, p 1-0-0; femora p 1-0-1, d 0-2-3, r 0-1-1. Epigyne (Fig. 
1 3B): similar toplatnicki and signatus, but readily separated by the comma-like ducts. 

Dimensions (mm): total length c. 10*0; carapace length 3*76, breadth 3*32, height 
2-36; abdomen length 5*92; eyes, anterior row 2*6, middle row 2*36, posterior row 
2-41; quadrangle length 2'04 (54% of carapace length). Ratios: AM : AL : 
PM : PL ::20 : 12 : 7'5 : 12; AL-PM-PL :: 11-5-1 7; AM : CL :: 20 : 5 (25% of AM diameter). 

MALE PARALECTOTYPE. In fair condition. Carapace (Fig. 13 A): amber with vague 
brownish markings; rubbed. Eyes: with black surrounds except AM; irregularly fringed by 
creamy white hairs. Clypeus: fringed in greyish white hairs. Chelicerae: elongate, porrect and 
diverging distally; amber, shiny; very sparsely clothed in light amber hairs; pro- and 
retromargins with three teeth. Maxillae and labium: yellow-brown. Sternum: pale 
yellow-brown with darker margins. Coxae: yellow-brown. Abdomen: yellow-brown with 



Fig. 13 Cocalodes longipes (Thorell). Paralectotype d 1 : A, dorsal; C, palp, retrolateral; E, 
median apophysis; G, palp, ventral. Lectotype 9: B, epigyne; D, carapace, lateral; F, dorsal. 

faint sooty lateral markings; spinnerets yellow-brown. Legs: generally orange-brown; spines 
strong and numerous. Spination of legs I: metatarsi v 2-2-2, p 1-1-1 , r 1-0-1 ; tibiae v 4^-0, 
p 0-1 -l,d 0-1-0, 1-1-1; patellae p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-2-0. Palp 
(Fig. 1 3C, E, G): the triangular tip of the functional conductor is distinctive. 

Dimensions (mm): total length c. 9*9; carapace length c. 4*28, breadth 3*64, height 
2*52; abdomen length 6 - 24; eyes, anterior row 2*88, middle row 2'55, posterior row 
2*68; quadrangle length 2-32 (54% of carapace length). Ratios: AM : AL : 
PM: PL:: 22-5: 14:8.5: 14:5; AL-PM-PL :: 13-17; AM : CL :: 22-5 : 8 (35% of AM 

286 F. R. WANLESS 

DISTRIBUTION. Indonesia: Ceram Island; Papua New Guinea: Yule Island. 

MATERIAL EXAMINED. Indonesia, Wahai, Ceram Island, paralectotype d 1 , 1872, (L. M. 
D'Albertis). Papua New Guinea, Yule Island (Roro), lectotype 9, 1872 (L. M. D'Albertis) 
(MCSN, Geneva). 

REMARKS. To judge from the geographical distribution, the male and female described above 
may not be conspecific. 

Coca lodes platnicki sp . n . 

DIAGNOSIS. From other species of Cocalodes by the combined absence of peg-like 
cymbial spines and curved tip of the functional conductor (Fig. 14F) in males. 
Females are readily separated by the absence of an epigynal septum and broad 
posterior ledge (Fig. 14H). 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 14B, C): yellow-brown with dark 
brownish sides and thin yellow-brown border line; pale areas including border line 
clothed in short whitish hairs, elsewhere covered in short blackish hairs. Eyes: with 
black surrounds except AM; fringed by whitish hairs. Clypeus: fringed in long white 
hairs. Chelicerae: moderately elongate, robust, inclined anteriorly and slightly divergent; 
pale orange-brown; sparsely fringed by long white hairs along outer basal margin; pro- and 
retromargins with three teeth. Maxillae and labium: yellow-brown. Sternum (Fig. 14G): 
yellow-brown with darker margins; shiny. Coxae: yellow-brown; shiny. Abdomen: with two 
pairs of impressed spots: pale yellow with blackish lateral bands; spinnerets pale yellow- 
brown tinged black. Legs: legs I amber grading to pale yellow-brown distally with blackish 
markings on femora, distal half of tibiae and metatarsi; legs II similar to I, but markings less 
distinct; legs III-IV yellow-brown with vague sooty annuli on metatarsi and darkening 
towards apices of tibiae; spines strong and numerous. Spination of legs I: metatarsi v 2-2-2, 
p 1-1-1 , r 1-0-1 ; tibiae v 2-2-2, p 1-1-2, d 0-0-1 , r 1-1-2; patellae p 1-0-0, r 1-0-0; femora 
p 2-1-2, d 0-3-2, r 1-2-2. Palp (Fig. 14D-F). 

Dimensions (mm): total length 7*2; carapace length 2*98, breadth 2-48, height 1'76; 
abdomen length 4*24; eyes, anterior row 2*28, middle row 2'02, posterior row 2' 12; 
quadrangle length 1-8 (60% of carapace length). Ratios: AM : AL : 
PM : PL :: 18 : 1 1 : 7 : 1 1-5; AL-PM-PL :: 10-14; AM : CL :: 18 : 4 (22% of AM diameter). 

FEMALE PARATYPE. In good condition. Carapace (Fig. 14 A): dark orange-brown with 
yellow-brown markings; dark areas clothed in short blackish hairs with whitish hairs 
elsewhere. Eyes: more or less as in d. Clypeus: fringed by white hairs. Chelicerae: 
robust and moderately porrect; amber; sparsely clothed in whitish and pale amber 
hairs; pro- and retro-margins with three teeth. Maxillae and labium: pale yellow- 
brown. Sternum: pale yellow-brown; thinly covered in testaceous hairs. Coxae: pale 
yellow-brown. Abdomen: similar to d, but ventrally with a pale greyish band from 
epigyne to spinnerets. Legs: legs I-II yellow-brown with vague darker markings and 
blackish femoral stripes; other legs yellow-brown with sooty spots on femora, darkened tibial 
apices and sooty annuli of metatarsi; spines strong and numerous. Spination of legs I: 
metatarsi v 2-2-2, p 1-0-0, d 0-1-2, r 1-0-0; tibiae v 2-2-2, p 1-1-1, d 0-1-0, r 1-1-1; 
patellae p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-2-0. Epigyne (Figs 14H-J; 21C-F): 
only species in genus known to have secondary spermathecae. 

Dimensions (mm): total length 8*96; carapace length 3-44, breadth 2-84, height 2'0; 
abdomen length 5*28; eyes, anterior row 2'52, middle row 2' 18, posterior row 2-28; 
quadrangle length 2*0 (58% of carapace length). Ratios: AM : AL : 
PM:PL:: 19 : 12 : 8 : 12; AL-PM-PL :: 13-16-5; AM : CL :: 19 : 4 (21% of AM diameter). 



Fig. 14 Cocalodes platnicki sp. n. Holotype cf: B, carapace, lateral; C, dorsal; D, palp, 
retrolateral; E, median apophysis; F, palp, ventral; G, sternum. Paratype 9: A, dorsal; 
H, epigyne; I, vulva, dorsal; J, vulva, ventral. 

VARIATION. Paratype cf measures 9*6 mm total length, 3 - 72 mm carapace length. Paratype 9 
c. 9-1 mm total length, 3'24 mm carapace length. 


MATERIAL EXAMINED. Irian Jaya, Sukarnapura (Hollandia): tropical rain forest, holotype d 1 , 
paratype 9, 31.xii.1944 (L. W. Saylor) (AMNH, New York); paratype 9, 5. i. 1945 (L. W. 
Saylor) (AMNH, New York); paratype rf, rain forest, xii.1944, (H. Hoogstraat) (AMNH, 
New York). 

ETYMOLOGY. This species is named after Dr N. I. Platnick, American Museum of 
Natural History, New York. 



Cocalodes in not a hi I is sp. n. 
(Figsl5A-F;18A, B) 

DIAGNOSIS. Similar to males of C. longipes and C. longicornis, but separated by the 
combined presence of peg-like cymbial spines and the finger-like dorsal prong of the 
median apophysis (Fig. 1 5D, E). 

FEMALE. Unknown. C. expers from Fergusson Island may belong here or possibly the 
female of C. longipes, from Yule Island (Papua New Guinea), which seems doubtfully 
matched with the male known only from Seram (Indonesia). 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 15 A, B): orange-brown with dark 
reddish brown sides and light orange margins; weakly iridescent under some angles of 
illumination; rubbed. Eyes: with black surrounds except AM; fringed in whitish and 
pale amber hairs. Clypeus: sparsely fringed by fine pale brownish hairs. Chelicerae: 
moderately long, inclined anteriorly; dark reddish brown, weakly iridescent blue under some 
angles of illumination; thinly covered in stiff brownish hairs; pro- and retromargins with 
three teeth; horn moderately long, but not protruding beyond clypeus. Maxillae: 
orange-brown to light yellowish along inner and distal margins. Labium: orange-brown 
tipped light yellow. Sternum: pale amber with darker margins; shiny. Coxae: pale amber. 
Abdomen: with two pairs of impressed spots; yellow-brown with brownish lateral bands 
above and central blackish band below; rubbed; spinnerets light orange-brown tinged black. 
Legs: tarsi light amber, metatarsi dark brown with light brownish hairs grading to amber with 

Fig. 15 Cocalodes innotabilis sp. n. Holotype rf: dorsal; B, carapace, lateral; D, palp, 
retrolateral; E, median apophysis; F, palp, ventral. Paratype cf: C, carapace, dorsal, 
showing cheliceral horn. 


white hairs proximally, tibiae dark brown, patellae brownish grading to amber proximally, 
femora amber; other legs amber; spines strong and numerous. Spination of legs I: metatarsi 
v 2-2-2, p 1-0-0, d 0-1-2, r 1-0-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0, r 1-1-1; patellae 
p 1-0-0, r 1-0-0; femora p 1-1-1, d 0-2-3, r 0-1-1. Palp (Figs 15D-F; 18A, B); the lobe 
associated with the tip of the functional conductor (arrowed, Fig. 15F) is also 
characteristic of this species. 

Dimensions (mm): total length 6'96; carapace length 2*88, breadth 2*4, height 1'84; 
abdomen length 4-0; eyes, anterior row 2*26, middle row 2*04, posterior row 2- 18; 
quadrangle length l - 72 (59% of carapace length). Ratios: AM:AL: 
PM : PL :: 18 : 1 1 : 6 : 1 1; AL-PM-PL :: 10-13; AM : CL :: 18 : 5 (27% of AM diameter). 

VARIATION, rf total length varies from 5'9 to 8 - 2 mm, carapace length 2'28-4 - 4 mm 
(eight specimens). 

In one male, having the same carapace length as the holotype, the cheliceral horn 
protrudes beyond the clypeus and may be clearly seen in dorsal view (Fig. 15C). Also, 
as in the case of C. papuanus the cymbium bears either two or three peg-like spines. 

DISTRIBUTION. Papua New Guinea. 

MATERIAL EXAMINED. Papua New Guinea: Louisiade Archipelago, Sudest Island, (L. 
J. Brass, Fifth Archbold Expedition to New Guinea) (AMNH, New York): Rambuso, 
on north coast, 0-100 m, holotype d 1 , camp 11, xiii.1956; Mt. Riu, paratype rf, 
250-350 m, 1956. Kokoda, paratype d 1 , viii.1933 (L. E. Cheesman) (BMNH. 
1934.12.14.269); Morobi Province, Buro river, paratype d, ix.1979 (/. H. Martin} 
(BMNH. 1981.7.31.1); Milne Bay, paratype cf, autumn 1944 (R. B. Burrows, A.P.O. 
928) (AMNH, New York); Oro Bay; paratypes 2c?d", (Lt. B. Struck, A.P.O. 503) 
(AMNH, New York), paratype d 1 , vii-viii. 1 943 (Sidney Sandier) (AMNH, New York). 

Genus ALLOCOCALODES gen. n. 

DEFINITION. Based on two species known only from males. 

Spiders of medium size (i.e. 4*0 to 8'0 mm in length). Male with elongate and 
porrect chelicerae bearing a median horn which is sometimes visible in dorsal view; 
not hirsute. Carapace: profile as in Figs 16 A, B; 17A, B); high, longer than broad, 
with steep thoracic slope, widest at level of coxae II-III; fovea long, sulciform, centre 
situated just behind posterior margin of posterior lateral eyes. Eyes: large with black 
surrounds except anterior medians; posterior median and posterior laterals set on 
pronounced tubercles; set in three transverse rows comprised of anterior medians 
(AM) and anterior laterals (AL), posterior medians (PM) and posterior laterals (PL); 
anteriors contiguous with apices procurved in frontal view and recurved in dorsal 
view; anterior medians largest; anterior laterals greater than half diameter of anterior 
medians; posterior medians large, positioned nearer to anterior laterals than to posterior 
laterals and more or less on optical axis of anterior laterals; posterior laterals about as large as 
anterior laterals and set inside lateral margins of carapace when viewed from above; 
quadrangle formed by posterior medians and posterior laterals broader than long and widest 
posteriorly; entire quadrangle, measured from between bases of anterior medians to posterior 
margins of posterior laterals, occupying between 60-65% of carapace length. Clypeus: 
between 25 and 37% of diameter of anterior median eyes. Chelicerae: elongate and porrect, 
sometimes diverging, with a median horn arising basally; promargin with four teeth, 
retromargin with two. Maxillae (Fig. 1 6C): long and diverging with rounded apices. Labium: 
oblong about half maxillae length. Sternum (Fig. 16C): scutiform. Coxae: I and II 
largest. Pedicel: short. Abdomen: elongate ovoid; spinnerets moderately long, posteriors 
longest with moderately long apical articles, anteriors robust, slightly longer than more 
slender medians; spiracle a transverse slit just in front of anterior spinnerets; tracheal system 
not examined; position of colulus indicated by scanty tuft of fine hairs; anal tubercle a broad 
cone. Legs: long and slender; spines moderately strong and numerous; claws pectinate; tufts 



present; scopulae absent. Male palps: moderately simple and similar in appearance; femora 
slightly bowed; patellae moderately long; tibiae with small dorsal tubercle, bifid retrolateral 
apophysis, the ventral element broad and rounded, the dorsal slender with translucent tip; 
cymbium with distal finger-like extension and shallow dorsally ridged excavation on retro- 
lateral basal margin; embolus (e) short, moderately stout and curved, arising subapically; 
functional conductor (c) a finger-like extension of the tegulum the tip of which lies either 
above or below that of the embolus; also arising from tegulum a lobe-like median apophysis 
(m); tegulum (t) ovoid with peripheral seminal ducts and what appears to be an inner folded 
duct. Expanded palps not examined. 

TYPE SPECIES. Allococalodes alticeps sp. n. 

AFFINITIES. Closely related to Cocalodes, see remarks on p. 264. 

DIAGNOSIS. From Cocalodes by the bifid retrolateral tibial apophysis, lobe-like functional 
conductor and median apophysis (Figs 16E; 17C). 

Fig. 16 Allococalodes alticeps sp. n. Holotype rf: A, dorsal; B, carapace lateral; C, 
sternum, coxae and mouth parts; D, palpal tibia, retrolateral; E, palp, ventral; F, palp, 


List of species in the genus Allococalodes gen. n. 

Allococalodes alticeps sp. n. 
A. cornutus sp. n. 

Allococalodes alticeps sp. n. 

(Fig. 16A-F) 

DIAGNOSIS. From A. cornutus by the pointed tip of the median apophysis (Fig. 16E). 
FEMALE. Unknown. 

MALE HOLOTYPE. In fair condition. Carapace (Fig. 16 A, B): dark chestnut brown with 
central yellow-brown thoracic band and vague yellow-brown markings in eye region; 
sides clothed in short recumbent amber hairs, iridescent under some angles of illumination, 
with whitish hairs on thoracic band and eye region. Eyes: with black surrounds except AM; 
fringed by whitish hairs. Clypeus: fringed in white hairs. Chelicerae: elongate and porrect; 
dark brownish orange with blackish mottling; thinly clothed in stiff light amber hairs; 
promargin with four teeth, retromargin with two; cheliceral horn protruding just beyond 
clypeal margin, but not visible in dorsal view. Maxillae and labium: brownish orange 
suffused black with inner margin of maxillae and labial tip dull yellow-brown. Sternum: 
brownish orange suffused black. Coxae: yellow-brown. Abdomen: mottled black with dorsal 
light yellow band somewhat broken and having blackish markings; clothed in minute dull 
whitish hairs; spinnerets tinged black, the posteriors with lighter terminal articles. Legs: 
generally yellow-brown with blackish annuli; spines strong and numerous. Spination 
of legs I: metatarsi v 2-0-1, p 1-1-1, r 1-1-1; tibiae v 2-2-2, p 1-1-1, d 1-1-0, 
r 1-1-1; patellae p 0-1-0, r 0-1-0; femora p 1-1-1, d 0-2-2, r 0-1-0. Palp (Fig. 
16D, E): when viewed from below (i.e. Fig. 16E), the tip of the embolus lies above the 
tip of the functional conductor (c.f. A. cornutus). 

Dimensions (mm): total length 4-84; carapace length 2*2, breadth 2*04, height 1'64; 
abdomen length 2-56; eyes, anterior row 1'96, middle row 1*68, posterior row 1-8; 
quadrangle length 1'44 (65% of carapace length). Ratios: AM:AL: 
PM : PL :: 16 : 10 : 6'5 : 10; AL-PM-PL :: 8-10; AM : CL :: 16 : 4 (25% of AM diameter). 


MATERIAL EXAMINED. Irian Jaya: Sukarnapura (Hollandia), rain forest, holotype d 1 , 
3 1 .xii. 1 944 (L. W. Saylor) (AMNH, New York). 

Allococalodes cornutus sp. n. 

(Fig. 17A-E) 

DIAGNOSIS. From A. alticeps by the obtuse tip of the median apophysis (Fig. 17C). 
FEMALE. Unknown. 

MALE HOLOTYPE. In poor condition. Carapace (Fig. 17A, B): dark orange-brown with 
pale eye region and median light yellowish brown thoracic band from foveal area to 
posterior margin; sides clothed in short recumbent amber hairs, shining under some 
angles of illumination, with dull whitish hairs on thoracic band and eye region. Eyes: 
with black surrounds except AM; fringed by dull whitish hairs. Clypeus: White 
haired. Chelicerae: elongate, porrect and diverging; dark orange-brown with blackish 
mottling; sparsely clothed in fine clear whitish hairs; promargin with four teeth, 
retromargin with two; cheliceral horn spiralled and protruding beyond clypeal margin. 
Maxillae: brown-black to dirty yellow-brown. Labium: dark brownish. Sternum: amber with 
irregular blackish mottling. Coxae: yellow-brown. Abdomen: damaged; elongate ovoid; 
yellow-brown with irregular blackish markings. Legs: generally light orange-brown grading 



Fig. 17 Allococalodes cornutus sp. n. Holotype d: A, carapace lateral; B, carapace, 
dorsal; C, palp, ventral; D, palpal tibia, retrolateral; E, palp, retrolateral. Abbreviations: c, 
functional conductor; e, embolus; m, median apophysis; t, tegulum. 

to yellow-brown distally with blackish annuli; spines moderately strong and numerous. 
Spination of legs I: metatarsi v 2-1-1, p 1-1-1, d 0-0-1, r 1-0-1; tibiae v 2-4-1, p 1-1-1, 
d 1-1-0, r 1-1-0; patellae p 1-0-0, r 1-0-0; femora p 1-1-0, d 0-2-2. Palp (Fig. 17C-E): 
when viewed from below the functional conductor lies above the embolic tip (Fig. 1 7C). 

Dimensions (mm): total length c. 4*8; carapace length 2-32, breadth 2*02, height 
l - 66; abdomen length c. 2*6; eyes, anterior row 1*94, middle row l - 62, posterior row 
1-76; quadrangle length 1-4 (60% of carapace length). Ratios: AM : AL : 
PM : PL :: 15: 9 : 6-5 : 9'5; AL-PM-PL :: 9-9; AM : CL :: 15 : 5'5 (37% of AM diameter). 


MATERIAL EXAMINED. Irian Jaya, holotype d 1 , (Dr. Moszkowski, no 54) (MNHU, Berlin. 


Taxonomic summary 

1 . Allococalodes gen. n., is proposed. 

2. Eight new species are described: 

Allococalodes alticeps sp. n. 
Allococalodes cornutus sp. n. 
Cocalodes cygnatus sp. n. 
Cocalodes expers sp. n. 
Cocalodes innotabilis sp. n. 
Cocalodes longicornis sp. n. 
Cocalodes platnicki sp. n. 
Cocalodes turgidus sp. n. 

3. Three species are newly synonymized: 

Cocalodes armatissimus Strand, 1913, and Cocalodes plebejus Szombathy, 1915, are 
junior synonyms of Cocalodes papuanus Simon, 1900. 

Cocalodes melanognathus Pocock, 1897, is a junior synonym of Cocalodes leptopus 
Pocock, 1897. 


I wish to thank the following colleagues for providing specimens for study. Dr Gianna 
Arbocco, Museo Civico di Storia Naturale, Geneva, Italy (MCSN, Genova); Dr M. 
GrasshofT, Forschungsinstitut Senckenberg, Natur-Museum, Senckenberg, Frankfurt am 
Main, West Germany (FS, Frankfurt am Main); Dr M Hubert, Museum National 
d'Histoire Naturelle, Paris, France (MNHN, Paris); Dr H. W. Levi, Museum of 
Comparative Zoology, Harvard, U.S.A. (MCZ, Harvard); Dr M. Moritz, Museum fur 
Naturkunde der Humboldt-Universitat, Berlin, East Germany (MNHU, Berlin); Dr 
N. I. Platnick, American Museum of Natural History, New York, U.S.A. (AMNH, 
New York). 

I also wish to thank Mr D. Macfarlane (CIE, London) for reading the manuscript 
and Mr P. York (BMNH. Photo unit) for the cleared epigyne photographs. 


Bonnet, P. 1956. Bibliographia Araneorum. 2 (2) : 919-1925. Imprimerie Douladoure, Toulouse. 

Forster, R. 1980. Unidentified objects. /. Arachnol. 8 (1) : 91-92. 

Neave, S. A. 1939. Nomencl. Zool. Vol I A-C: 957 pp. Zoological Society, London. 

Petrunkevitch, A. 1928. Systema Aranearum. Trans. Conn. Acad. Arts Sci. 29 : 270 pp. 

Platnick, N. I. & Shadab, M. U. 1975. A revision of the spider genus Gnaphosa (Araneae: 

Gnaphosidae) in America. Bull. Am. Mus. nat. Hist. 155 : 3-66. 
Pocock, R. I. 1897. Spinnen (Araneae). In Kiikenthal, W., Ergebnisse einer zoologische 

Forschungsreise in dem Molukken und Borneo. Abh. senckenb. naturforsch. Ges. 23 : 591-629. 
Prdszyriski, J. 1971. Catalogue of Salticidae (Aranei) specimens kept in major collections of 

the world. Annls zool. Warsz. 28 : 367-5 19. 
Roewer, C. F. 1954. Katalog der Araneae. 2 Abt. B: 924-1290. Institut Royal des Sciences 

Naturelle de Belgique, Bruxelles. 
Simon, E. 1900. Etudes arachnologiques 30e Memoire (1) XLVII Descriptions d'especes nouvelles de 

la famille des Attidae. Annls Soc. ent. Fr. 69 : 27-6 1 . 
1901. Histoire Naturelle des Araignees. 2 (3) : 381-668. Roret: Libraire Encyclopedique, 

Strand, E. 1913. Neue indoaustralische und polynesische Spinnen des senckenbergischen Museums. 

Arch, naturgesch. 79A (6) : 1 13-123. 
1915. Indoaustralische, papuanische und polynesische Spinnen des senckenbergischen Museums, 

gesammelt von Dr E. Wolf, Dr J. Elbert u. a. In wissenschaftliche Ergebnisse de Hanseatischen 

Siidsee Expedition 1909. Abh. senckenb. naturforsch. Ges. 36 (2) : 181-274. 

294 F. R. WANLESS 

Szombathy, K. 1915. Attides nouveaux appartenant aux collections du Musee National Hongrois. 

Annls hist.-nat. Mus. natn. hung. 13 : 468-490. 
Thorell, T. 1878. Studi sui ragni Malesi e Papuani. Part II. Ragni di Amboina raccolti da Prof. 

O. Beccari. Mus. civ. Stor. nat. Giacomo Doria 13:317 pp. 

1881. Studi sui ragni Malesi e Papuani. Part III. Ragni dell Austro-Malesia e del Capo 

York, conservati nel Museo Civico di Storia Naturale di Geneva. Mus. civ. Stor. nat. 
Giacomo Doria 17 : 720 pp. 

Wanless, F. R. 1978. A revision of the spider genera Belippo and Myrmarachne (Araneae: 
Salticidae) in the Ethiopian region. Bull. Br. Mus. nat. Hist. (Zool.)33 (1) : 139 pp. 

1981. A revision of the spider genus Cocalus (Araneae: Salticidae). Bull. Br. Mus. nat. 

Hist.(Zool) 41 (5): 253-261. 

Waterhouse, C. O. 1 902. Index zoologicus 42 1 pp. London. 

Manuscript accepted for publication 5 October 198 1 



Fig. IS (A, B) Cocalodes innotabilis sp. n. cT palp: A, peg-like cymbial spines, x500; B, 
median apophysis and functional conductor, xlOO. (C, E) C. papuanus Simon, rf palp: C, 
median apophysis, xlOO; E, ventrolateral view, x60. (D) C. thoracicus Szombathy. d 
palp, median apophysis, xl 50. 




Fig. 19 (A, B) Cocalodes papuanus Simon. Vulva: A, ventral; B, dorsal, xlOO. (C, D) C. 
leptopus Pocock. Vulva: C, ventral; D, dorsal, x!40. (E, F) C. cygnatus sp. n. Vulva: E, 
ventral; F, dorsal; x 140. 



Fig. 20 (A, B) Cocalodes macellus (Thorell). Vulva: A, ventral; B, dorsal; xlOO. (C, D) C. 
thoracicus Szombathy. Vulva: C, ventral; D, dorsal; xlOO. (E, F) C. turgidus sp. n. 
Vulva: E, ventral, xlOO; F, dorsal, x!35. 




Fig. 21 A, Cocalodes longicornis sp. n., vulva dorsal view, x!40. B, C. papuanus Simon, 
cheliceral horn, x85. (C, D) C. platnicki sp. n. Vulva: C, ventral; D, dorsal; xlOO. (E, F) 
spherical bodies in vulva of C. platnicki; E, x280; F, x!200. 

Anatomy and evolution of the jaws in the 
semiplotine carps with a review of the Genus 
Cyprinion Heckel, 1843 (Teleostei: Cyprinidae) 

Gordon Howes 

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 


Scant attention has been paid to the jaw anatomy of cyprinid fishes. Takahasi (1925) 
described the jaw musculature of various cyprinoids, but only Matthes (1963) and Alexander 
(1966) have described the osteo-myological anatomy of the cyprinid jaw. These authors, and 
Ballantijn (1969), have also attempted a functional interpretation of particular jaw types. 
Howes (1978, 1979, 1980, 1981) described and commented on jaw types of (mostly) non- 
barbelled carps ('Leuciscinae'). But, in these studies the potential value of the jaws as 
indicators of phyletic relationship was not recognised. The more diverse jaw morphology of 
barbelled carps ('Cyprininae') has, however, focussed attention on this aspect of jaw 

From among the 'Cyprininae' the genus Semiplotus is chosen as a model for detailed 
description because member species possess what is an obviously highly derived osteo- 
myological jaw structure, manifest externally in the size and shape of the mouth (see below). 
Semiplotus species are large (according to Day (1889) S. semiplotus attains at least 2 feet in 
length), distinctive carps whose geographical range extends through northern India and 
Burma. The distinctiveness of these fishes lies in the size and shape of the head and extreme 
body depth (Figs 1A-C). Likewise, the mouth is of great width (probably the widest mouth 
in relation to head length of any cyprinid fish), the lower jaw is edged with a cornified 
epithelium and is overhung by the upper jaw (Figs 1 B & C). 

Many cyprinid taxa possess broad, ventral mouths with cornified cutting edges, the 
so-called 'sector mouth' (see p. 313 for definition). Only in Onychostoma, Varicorhinus, 
Capoeta and Cyprinion does the morphology of the mouth approach that of Semiplotus, 
both in width and complex development of the lower lip. 

Members of these genera are compared here in order to test the hypothesis that a sector 
mouth is synapomorphic for the group and also to elucidate the possible evolutionary 
pattern of semiplotine jaw modification. 

Nomenclatural note 

In the anatomical descriptions, Scaphiodonichthys burmanicus is referred to as Semiplotus 
burmanicus. The existence in both 'genera' of synapomorphic osteological and myological 
characters indicates their generic unity (see p. 312 for a list of characters, and p. 331 for 
complete synonymy). No specimens of Semiplotus modestus have been examined and 
reference to 'Semiplotus' in the anatomical text refers only to the species semiplotus and 

List of specimens used in anatomical description 

Capoeta capoeta BMNH 1968.12.13 : 405^16 

Capoeta capoeta BMNH 1879.1 1.14 : 19 (skeleton) 

Bull. Br. Mm. not. Hist. (Zool.)42 (4) : 299-335 Issued 24 June 1982 



Fig. 1 Semiplotus semiplotus. A, in lateral view. B & C, ventral and anterior aspects of the head. 
Scale = 10 mm. Drawn from specimen 1889.2.1 :365-6. 

Cyprinion acinaces 
Cyprinion kais 
Cyprinion macrostoma 
Cyprinion microphthalmum 
Cyprinion microphthalmum 
Cyprinion milesi 
Cyprinion watsoni 
Cyprinion watsoni 
Onychostoma laticeps 
Onychostoma varus 
Semiplotus burmanicus 
Semiplotus burmanicus 
Semiplotus burmanicus 
Semiplotus burmanicus 
Semiplotus semiplotus 
Semiplotus semiplotus 
Semiplotus semiplotus 
Squaliobarbus curriculus 

BMNH 1899.7. 15 26 

BMNH 1977.8.25 1-5 

BMNH 1974.2.22 1115-8 

BMNH 1974.2.22 118-95 

BMNH 1883.8.2:4-9 

BMNH 1977. 12. 13: 532-545; 546-556 

BMNH 1889.2.1 : 263^ 

BMNH 1889.2.1 : 387-396 

BMNH 1913.4.15:90-99 

BMNH 1969.4.15: 105-110 

BMNH 1935.4.18:22 

BMNH 1893.2.16 : 32 (skeleton) 

BMNH 1889. 10.4: 2 

BMNH 1920.9.8: 1-3 

Uncat. Smithsonian Colin. 

BMNH 1889.2.1 : 365-6 

BMNH 1889.2.1 : 367 (skeleton) 

BMNH 1972.7.26: 10 

BMNH 1 888.5. 1 5 : 29 (skeleton) 


Varicorhinus beso BMNH 1968.7.24 : 2 

Varicorhinus beso BMNH 1968.7.24 : 17-19 (alizarins) 

Varicorhinus beso BMNH 1902.12.13 : 377 (skeleton) 

Varicorhinus tanganicae BMNH 1906.9.6 : 1 1 (skeleton) 

Abbreviations used in text figures 

A ( a, A,D, A 2 , A 3 , 

A w Divisions of the adductor mandibulae muscle 

Aa Anguloarticular 

Aamp Aponeurosis of adductor mandibulae and pro- 
tractor hyoideus muscles 

Aap Premaxillary anterior ascending process 

Afl, 2, 3 Articulatory facets of anguloarticular 

Ahy Anterohyal 

Ama Aponeurosis of adductor mandibulae A 1 muscle 

Ao Adductor operculi muscle 

Apl, 2 Articulatory processes of anguloarticular 

Br Branchiostegal ray (numbered) 

Cm Coronomeckelian bone 

Cp Coronoid process of dentary 

Ctls Labial shelf connective tissue 

Cts-fef Connective tissue strand linking maxilla to frontal- 
ethmoid foramen 

De Dentary 

Dhy Dorsohyal 

Do Dilatator operculi muscle 

Ect Ectopterygoid 

Elt Epithelial layer of lower lip 

Ent Entopterygoid 

Epo Epioccipital 

Etb Epithelial tendinous layer 

Exo Exoccipital 

Fr Frontal 

Fsef Frontal-supraethmoid foramen 

Hy Hyomandibula 

Ic Intercalar 

Ihy Interhyal 

Im Intermandibularis muscle 

lop Interoperculum 

lopf Interopercular facet 

Ke Kinethmoid 

Ks Keratinized lip surface 

Lac Ligament linking retroarticular with anterohyal 

Lai Ligament linking retroarticular with 


Lap Levator arcus palatini muscle 

Lapo Ligament linking retroarticular with preoperculum 

Laq Ligament linking anguloarticular with quadrate 

Le Lateral ethmoid 

Lkee Ligament linking kinethmoid to ethmoid 

Lkmx Ligament linking kinethmoid to maxilla 

302 G. J. HOWES 

Lkp Ligament linking kinethmoid to palatine 

Lkpm Ligament connecting kinethmoid to premaxilla 

LI Lower lip 

Llm Lateral lamellae 

Lo Levator operculi muscle 

Lpe Ligament linking palatine to ethmoid 

Lpm Ligament linking palatine to maxilla 

Lqu Ligament linking quadrate to interoperculum 

Ls Labial shelf 

Mdp Maxillary distal process 

Me Mesethmoid 

Met Metapterygoid 

Mfp Medial fatty partition 

Mkl Medial keratinized layer of lower lip 

Mlp Maxillary mid-lateral ascending process 

Mlt Medial tissue layer of lower lip 

Mmp Medial maxillary process 

Mn Medial notch 

Mv Maxillary valve 

MX Maxillary 

Mxf Maxillary foramen 

Mxn Maxillary notch 

Nc Neural complex 

Ns Neural spines 

Op Operculum 

Pa Parietal 

Pal Palatine 

Pe Preethmoid 

Ph Protractor hyoideus muscle 

Pha Medial aponeurosis of protractor hyoideus 

Phat Anterior tendinous segment of protractor hyoideus 

Phav Antero- ventral segment of protractor hyoideus 

Phbt Basal tendinous layer of protractor hyoideus 

Phd Dorsal segment of protractor hyoideus 

Phi Lateral segment of protractor hyoideus 

Phlt Lateral tendon of protractor hyoideus 

Phpv Postero- ventral segment of protractor hyoideus 

Phy Posterohyal 

Pop Preoperculum 

Pro Prootic 

Ps Parasphenoid 

Pte Pterotic 

Pty Pterygiophore (dorsal) 

Q Quadrate 

Qaf Quadrate anterior facet 

Qlf Quadrate lateral facet 

Qmb Quadrate medial buttress 

Ra Retroarticular 

Se Supraethmoid 

Sh Sternohyoideus muscle 

Sj Synarthritic mandibular joint 

Sn Supraneurals (numbered) 

Soc Supraoccipital 

Socr Supraoccipital crest 



Subtemporal fossa 


Symplectic facet 

Tendon connecting A 2 with A,b 

Insertion tendon of adductor mandibulae A 3 



Ventral epithelial lip tissue 


Ventral medial tissue layer of lower lip 



IP , Mn Mdp 

Fig. 2 Semiplotus semiplotus, upper jaw bones. A, premaxilla, lateral view. B, maxilla, lateral 
view. C, maxilla, medial view of symphysial portion; dashed line indicates meniscus. D, maxilla, 
dorsal view. Scale = 10 mm. 

Jaw anatomy of Semiplotus semiplotus (McClelland, 1839) 

Upper jaw bones 

The premaxilla (Fig. 2A) is exceedingly thin and shallow, with only a slight anterior 
ascending process. The posterior portion of the bone is directed ventrally at a right angle to 
its anterior part, with its tip compressed and shallowly bifurcated. 

The maxilla (Fig. 2B) is deep, its dorsal border convex. The anterior ascending process 
(Aap) is shallow and thick, its dorsal border notched (the palatine ligament lies across this 
notch). The upper symphysial border of the ascending process is turned so that its face forms 
a 45 angle to the midline. The lower portion is strongly concave and extends antero- 
medially to form the symphysial part to the maxilla, syndesmotically contacting its partner. 
The medial (rostral) maxillary process (Mmp, Fig. 2C) is almost square and it too makes 

304 G. J. HOWES 

syndesmotic contact with its partner. A mid-lateral ascending process is represented by a 
slight convexity of the maxillary dorsal border (Mlp). This part of the maxilla is separated 
from the posterior part by a medial notch (Mn). The posterior rim of the notch acts also as 
the leading border of a broad medial triangular ledge. 







Fig. 3 Semiplotus semiplotus, upper jaw articulation in oblique dorso-lateral aspect. 

Scale = 5 mm. 

Articulation of the upper jaw (Fig. 3) 

The upper jaw bones articulate with the cranium via the kinethmoid, the palatine and the 

premaxillary meniscus. 

The kinethmoid (Ke) is attached via a thick ligament from its head to the ascending 
process of the premaxilla (Lkpm). A broad, bifurcated ligament (Lkee) extends from the 
posterior face of the kinethmoid to attach to the lateral edges of the supraethmoid and, 
dorsally, to the connective tissue overlying the ethmoid region. A strand of this connective 
tissue (Cts-fef) inserts onto the rim of the foramen between the frontals and the 
supraethmoid. A mid-lateral ligament (Lkp) attaches the kinethmoid to the palatine, and a 
ventro-lateral ligament (Lkmx) links it to the maxilla. 

A paired ligament (Lpm) attaches the lateral face of the maxilla with the lateral shelf of the 
palatine (Pal). The palatine, in turn, is connected with the mesethmoid by a thick medially 
directed ligament (Lpe). 

Interposed between the medial face of the maxilla, the preethmoid and part of the vomer is 
a cartilaginous meniscus (sensu Alexander, 1966). This is thick and ellipsoidal and lies 
between a ventral prominence of the premaxillary ascending process and the medial (rostral) 
process (Fig. 3). 

Muscles attaching to the upper jaw (Fig. 4) 

The adductor mandibulae A, muscle occupies only the lower half of the cheek, the area 
below the orbit and between it being filled with fatty connective tissue. The muscle is 
divisible into ventral and dorsal sections. The lower section, A,b, originates from the lower 
border of the preoperculum, is thin and triangular, but with its ventral border thickly 
tendinous. Fibre alignment varies from 45 (ventrally) to almost horizontal (dorsally). At the 
posterior tip of the maxilla, the muscle section is constricted into a bifurcated tendinous 







Fig. 4 Semiplotus semiplotus. A, lateral view of superficial jaw and cranial musculature. B, detail 

of jaw muscle insertions. Scale = 5 mm. 

area, the outer tendon inserting on a small lateral maxillary process, the inner on the medial 
maxillary notch (see above, p. 304). The lateral border of the inner tendon (Fmd, Fig. 4B) 
extends mesially to attach to the lateral face of the dentary coronoid process. The medial 
fascia of adductor A,b is connected with the lateral body of adductor mandibulae muscle A 2 
by a thick tendon (see below). 

The dorsal section of the adductor mandibulae, A,a, originates from the central area of the 
preoperculum. This portion of the muscle is even thinner than A,b, being only a few fibres in 
thickness at its origin. The segment is oblong, with an almost parallel arrangement of fibres. 
The muscle terminates via a narrow tendon which forms the lateral segment of a broad, thick 
aponeurosis (see below). The tendon of insertion reaches the first of the palatine-maxillary 

The aponeurosis into which muscle A,a inserts is complex (Ama, Fig. 4). Laterally it is 
joined to the medial face of the 1st infraorbital, and medially to the lateral ethmoid and to 
the palatines. Ventro-medially the sheet has a posterior extension in the form of a tendinous 
band which joins onto the anterior edge of the entopterygoid and then expands to become 
continuous with the medial fascia of muscle A 3 (Fig. 8A). 

Upper lip and buccal cavity (Fig. 9) 

Epithelial tissue covering the upper jaw is thin and lacks a cornified edge. The inside of the 
mouth is bordered by a single row of leaf-like lamellae; above this row is a thin, narrow 
maxillary valve (Mv). The tissue covering the antero-lateral buccal roof is formed into a near 
longitudinal series of broad leaf-like lamellae (Llm). The surfaces of these lamellae are highly 
papillose. The vomerine tissue is transversly rugose; there is no vomero-palatine organ 
(sensu Matthes, 1963). 





Fig. 5 Semiplotus semiplotus, lower jaw bones. A, lateral; C, dorsal; and D, posterior views. B, 
medial aspect of dentary joint. Dotted lines indicate the course of a ligamentous connection. 
Scale = 10 mm. 

Lower jaw bones (Fig. 5) 

Each dentary (Figs 5 A & B) articulates with its partner via a complex symphysial joint (Sj). 
This is basically a peg and socket joint, the central peg being on the dentary of the right side 
and inserting into a deep notch in the left dentary. Above the joint there is a strong ascending 
process on each bone, joined syndesmotically to that of its partner. Ventrally, the 
articulation is more complex (Fig. 5B). Each dentary is extended as a ventral process which is 
joined to its counterpart via a bifurcated, curved articulation. Strong, vertical ligaments 
connect each facet of this joint. 

The wall of the dentary is thick, its dorsal part tapering to a thin edge and curving mesially 
so as to form a sloped labial surface. On the medial underside of this lip is a deep cavity into 
which the intermandibularis muscle inserts. The lateral face of the dentary bears a deep ridge 
which tapers posteriorly to a V-shaped process. The coronoid process is high and almost 
square, its anterior edge curved outward so that when viewed dorsally (Fig. 5C) the plane of 
the dorsal margin of the coronoid is coincident with that of the body axis. The posterior 
border of the dentary is blunt with a rounded ventral process corresponding in position to the 
coronoid process (Dvp, Fig. 5 A). 

The anguloarticular (Aa, Figs 5C & D) is a large, triangular bone, its posterior dorsal edge 
extending someway up the medial face of the dentary coronoid process, following that 
processes' outward curvature. The articulatory surface is complex. Postero-laterally there is 
a triangular facet (Afl, Fig. 5D), its posterior border extending only slightly beyond that of 
the dentary. The medially directed portion of the anguloarticular is bifurcated. The dorsal 
process (Apl) is thickly rounded and directed somewhat dorsally, its posterior face forming a 
facet (Af2). The ventral process (Ap2) is triangular and is directed slightly ventrally. The 
medial dorsal process (Apl) is separated from the outer wall of the bone by a deep, concave 
channel. The posterior aspect of the anguloarticular is V-shaped and there is a large, figure of 
8-shaped facet situated between the dorsal and ventral medial processes (Af3). In lateral 



view, only a narrow portion of the anguloarticular is visible beyond the posterior dentary 

The retroarticular (Ra, Fig. 5D) is a thick wedge lying below the ventral anguloarticular 
process (Ap2). Laterally, the retroarticular is almost entirely covered by the ventral process 
of the dentary and only a narrow portion is visible (Fig. 5 A). 

The coronomeckelian (Cm, Fig. 5C) is an almost oblong bone with a rounded posterior 
border. Its anterior portion projects medially to form a deep, sloped shelf. There is a slight 
depression on the shelf into which insert the tendons of the adductor mandibulae 

Articulation of the lower jaw (Fig,. 6) 

The complex articulatory surface of the anguloarticular is matched by an equally complex 

quadrate surface. 

The quadrate facet occupies the lateral anterior and medial portions of the bone (Figs 6B & 
C). The lateral facet (Lqf) is aligned at 45 to the horizontal and apposes the medial face of 
the lateral anguloarticular process (Afl). The anterior facet is also aligned at 45 and apposes 

Ect i 





Fig. 6 Semiplotus semiplotus, lower jaw articulation. A, medial view of posterior jaw elements 
and their connections with suspensorial and opercular bones. B, anterior and C, lateral aspects of 
quadrate and inferior opercular elements. Scale = 5 mm. 



the central anguloarticular facet (Af3). The medial part of the quadrate is formed into a 
buttress with an elliptical anterior face which apposes the dorso-medial anguloarticular facet 
(Af2). The ventro-medial process of the anguloarticular (Ap2) lies below the medial quadrate 
buttress and is attached to it by a thick ligament (Laq). 

Ventrally a complexly divided ligament attaches the jaw to certain opercular and hyal 
elements. Immediately from its point of attachment to the posterior face of the retroarticular 
the ligament trifurcates and attaches, (1) to the anterior face of the interoperculum (Laio) 
which is medially curved and formed into a facet; (2) to the medial face of the preoperculum 
(Lapo) which is formed into a condyle; and (3) to the posterohyal via a fascial attachment to 
the ceratohyal (Lac). 

The lateral articulations between the anguloarticular, preoperculum and interopercular 
bones is covered by a broad ligament which stems from the ventro-lateral face of the 
quadrate (Lqi, Fig. 8A). 

Muscles attaching to the lower jaw (Fig. 7) 

The muscles having their attachment to the lower jaw are the adductor mandibulae A 2 and 

A 3 (Fig. 7B). The upper portion of muscle A 2 originates from the hyomandibula and the 



Ph B 

Fig. 7 Semiplotus semiplotus, lower jaw musculature. A, lateral view of the deeper muscle 
elements. B, medial view of jaw muscle insertions and their connection with hyoid musculature. 
Scale = 5 mm. 



lower lateral portion of the preoperculum. All its fibres insert into a medial aponeurosis. 
From the lateral part of this aponeurosis a narrow tendon runs antero-ventrally into the 
medial fascia of the adductor muscle (A,b (Fig. 7A). A bundle of fibres continues forward 
from the aponeurosis and almost immediately inserts into a thick tendon which terminates 
on the coronomeckelian bone (sesamoid articular). Insertion of A 2 is via a broad tendon onto 
the dorso-posterior rim of the dentary coronoid process. A thin, inner branch of the tendon 
continues ventrally down the medial face of the coronoid process to insert on the 
coronomeckelian bone. Attaching to this tendon is a triangular sheet of muscle fibres 
crossing from A 3 ; this most probably represents section A w of the adductor mandibulae 
series (see Fig. 7B & p. 3 1 3). 




Fig. 8. Semiplotus semiplotus, hyoid musculature. A, ventral aspect of head with right side 
dissected to display major hyoid muscle elements. Dotted lines indicate path of the inter- 
mandibularis muscle. B, detailed view of protractor hyoideus insertions on hyoid elements of the 
right side. Scale = 5 mm. 

310 G.J.HOWES 

Adductor mandibulae muscle A 3 originates partly from the hyomandibula (where it is 
separated from A 2 by the levator arcus palatini muscle) and partly from the deeply concave 
lateral face of the metapterygoid. Its fibres run almost at right angles to the midline to join 
the aponeurosis of muscle A 2 . Antero-ventrally, A 3 inserts via a strong tendon which runs 
through the deep cleft between the medial and lateral portions of the anguloarticular, to join 
an aponeurosis on the coronomeckelian bone (Fig. 7B). Extending anteriorly from this 
aponeurosis is a set of thin, tendinous fibres which spread along the inner aspect of the 
dentary upper margin. These tendinous bands are connected by fibrous tissue to the 
overlying loose, fatty connective tissue of the lower lip (see below). From the lateral part of 
the aponeurosis a tendon extends ventro-posteriorly, across the medial face of the angulo- 
articular, to join the lateral portion of the protractor hyoideus muscle (Tcph, Fig. 7B). 

The protractor hyoideus muscle is a short and thick element divisible into dorsal and 
ventral sections (Fig. 8A). The ventral portion is X-shaped, the two anterior arms separated 
by a thick nodule of fatty tissue and each attaching to the medial face of the dentary. Their 
posterior junction is a thick pad (Pha) from which radiate wide, tendinous bands to form the 
ventral face of the muscle. These bands insert into the posterior margin of the tissue forming 
the lower lip (see below). The posterior arms of the muscle are almost entirely tendinous and 
attach to the branchiostegal rays, and the inner faces of the posterohyal and the interhyal. 
Each dorsal section of the protractor hyoideus is broad and extends from the ceratohyal and 
the 1st branchiostegal ray to join the ventrohyal. The more medial fibres run into the body of 
the muscle and insert into the tendinous area just anterior to the ventrohyal (Fig. 8B). 

The intermandibularis is a thin, cylindrical muscle crossing between the dentaries where it 
is contained in a small cavity. The muscle separates, transversly, the anterior portions of the 
protractor hyoideus (Figs 8B & 9B). 

The lower lip (Fig. 9) 

The lower lip of Semiplotus semiplotus is a complex structure composed of several tissue 
types which extend well beyond the outer face of the dentary. Its core is a deep shelf of dense 
collagenous tissue that surrounds the dentary (Ls). This labial shelf is enveloped in loose, 
stratified epithelium. The tissue that covers the dorsal surface of the labial shelf is comprised 
of a thick, fatty epidermal layer (Elt), and an underlying, more fibrous tissue (Mlt). The 
ventral surface of this tissue adheres to tendinous bands originating from the aponeurosis 
that incorporates the insertions of the adductor mandibulae and protractor hyoideus muscles 
(see above). Thus, toward the lateral part of each dentary the labial epidermis becomes 
united with jaw and hyoid tendinous elements and only at the symphysis is it firmly attached 
to each dentary. Anteriorly, where it passes around the leading edge of the labial shelf, the 
epidermis is thin and cornified, forming a chisel-edge to the jaw (Ks). Ventrally, the 
epidermis is attached to the labial shelf by thin strands of connective tissue. As it passes 
beneath the dentary the tissue becomes tendinous and contiguous with the ventral face of the 
protractor hyoideus (see above). 

Apomorphic jaw characters in Semiplotus 

Before considering whether, or to what degree, the jaws of Semiplotus semiplotus may be a 
derived feature, it is necessary to present a model of the plesiomorph cyprinid jaw and its 
associated articular elements for comparison. The following model is modified from Howes 
(1980, 1981) and is reckoned to be the plesiomorph type on grounds of its wide-spread 
occurrence amongst various cyprinid taxa. 

The plesiomorph maxilla has a well-developed mid-lateral ascending process with convex 
anterior and concave posterior borders; the premaxilla L-shaped with a moderately 
developed anterior ascending process; dentary with a vertical, high, long coronoid process 
situated posteriorly; the anguloarticular with its dorsal border confluent with that of the 
dentary coronoid process but not extending to cover the medial face of that process, its 






Fig. 9 Semiplotus semiplotus. A, ventral view of upper lip and buccal cavity. B, sagittal section 
through the lower lip and associated hyoid musculature. Scale = 5 mm. 

posterior border reaching to well beyond the posterior margin of the dentary, its articulatory 
facet transversly concave. All those features are found in the jaws ofSqualiobarbus, Fig. 10. 

The plesiomorph cyprinid quadrate is regarded as one essentially broadly triangular in 
outline and with an anteriorly positioned articulatory facet. 

The plesiomorphic condition of the adductor mandibulae A, muscle in cyprinids would 
appear an undivided element having its insertion on the outer, posterior portion of the 
maxilla (as in aspinine cyprinids; see Howes, 1979). Commonly in cyprinids there are two 
medial sections of the adductor mandibulae. The outer (A 2 ) inserting on the rim of the 
coronoid process of the dentary, and the inner (A 3 ) inserting on the coronomeckelian bone 
(see Takahasi, 1925 : 20). Primitively, the mentalis section (A w ) of the adductor complex is 
present as a well-developed element (as in the bariliines, chelines, aspinines and cultrines) 
but in the majority of cyprinid taxa it is reduced or lacking entirely (see Takahasi, 1925 : 20). 

According to Takahasi (1925) and Matthes (1963) the protractor hyoideus ( = genio- 
hyoideus) muscle is usually attached to the 1st and/or 2nd branchiostegal rays, rarely to the 




Fig. 10 Squaliobarbus curriculus, jaw bones. A, maxilla. B & C, dentary in lateral and dorsal 

views. Scale = 5 mm. 

3rd and never to the interhyal. A tendinous lateral border of the muscle is not uncommon in 
cyprinids and in many taxa the lateral part of the muscle appears as a separate element. 

Semiplotus semiplotus and S. burmanicus show markedly derived states of the above 
hypothesized plesiomorph jaw and muscle morphology. The following features uniquely 
characterize the jaws of Semiplotus: 

Upper jaw 

1. Premaxilla: shallow, thin, with acute posterior curvature and reduced ascending 

2. Maxilla: deep, with broad anterior ascending process, reduced mid-lateral process and 
depressed, notched posterior portion. 

3. Bifurcated tendon of muscle A, attaching to lateral and medial faces of the maxilla. 
Some amplification of this character is required. It was noted above that the plesio- 
morph condition is for muscle A, to insert on the outer face of the maxilla. Most often 
this insertion is tendinous but in bariliine, cheline and aspinine cyprinids it may be 
musculose across a broad fascia that cover both the dorsal rim of the dentary as well as 
its lateral aspect (see Howes, 1976 Fig. 23). This, often tendinous, fascia may represent 
the 'ligamentum primordium' as might the connective tissue in other cyprinids which 
links the maxilla to the face of the anguloarticular and enwraps the insertion area of the 
lower jaw musculature. A divided tendinous insertion of the superficial adductor 
muscle is present in Cyprinus, Carassius and Gobio. However, unlike the semiplotine 


condition, the tendons cross over so that the insertion of the lower element (A,b) is 
above that of the upper (A,a). Takahasi (1925) regarded Cyprinus and Carassius as 
representing Transitional stages . . .' between the simple bariliine (Opsariichthys) and 
complex gobioine (Gobio) types of insertion. Crossing of A, tendons also occurs in 
Aspidoparia, a genus with bariliine affinities (see Howes, 1980), Cosmocheilus, 
Cyclocheilichthys, Puntioplites and Amblyrhynchichthys. Whilst it is possible that these 
four latter genera are closely related, such a relationship appears unlikely between them 
and Aspidoparia. All five taxa have a short, deep ethmoid region and the mouth is 
inferior in position. It is likely that the crossed tendon arrangement is functionally more 
efficient, in activating jaws of this kind and has been derived independently in the 
various taxa. 

4. Kinethmoid attached to connective tissue overlying the ethmoid bloc. 

Lower jaw 

5. Dentary: with a sloped labial surface, lateral ridge, outwardly curved coronoid process, 
ventral posterior process and a synarthritic joint. 

6. Anguloarticular rising dorsally to cover the medial face of the dentary coronoid 
process, only a small portion of the bone extending beyond the posterior border of the 
dentary; the medial portion extended and formed into stout and complex articular 

7. A medial tendon of muscle A 2 to which is attached muscle A w . Some amplification of 
this character is required: As noted above, the common, and presumed plesiomorph 
condition in cyprinids is for muscle A 2 to insert on the rim of the coronoid process of 
the dentary. In Semiplotus and some Cyprinion species (see p. 309) the muscle is 
bifurcated (see above) with both an outer tendinous insertion on the coronoid process 
and an inner one on the coronomeckelian bone. A segment of muscle stretches from the 
lower part of the tendon of muscle A 3 to the anguloarticular and should almost 
certainly be identified as an A w . Although a similarly developed muscle stems from the 
tendon of A 3 in Varicorhinus beso (p. 317) and some Barbus species (Banister, pers. 
comm.) it does not insert on a medial tendon of A 2 but on the medial face of the 

8. Protractor hyoideus muscle connected anteriorly, via an aponeurosis, with the adductor 
mandibulae insertions; its posterior part tendinous, attaching to all branchiostegal rays, 
posterohyal and interhyal. 

9. Lateral quadrate facet apposing lateral anguloarticulatory process. Quadrate short and 

Comparisons of the jaws in Semiplotus with those in Onychostoma, Varicorhinus, 

Capoeta and Cyprinion 

Taxa with 'sector mouths', i.e. an inferior mouth which is wide and has an exposed cornified 
mandibular cutting edge; the dentary invariably has a broad, sloping or deflected labial 
surface, occur frequently amongst the Cyprinidae. Examples are Onychostoma, 
Varicorhinus, Capoeta, Cyprinion, Barbus and some species of Schizothorax. Although a 
cornified lower jaw occurs in Labeo and Garra it is invariably covered by a thick 
labial fold. 

The jaw morphology of Onychostoma, Varicorhinus, Capoeta and Cyprinion species is 
remarkably similar to that of Semiplotus. The similarities may be independently derived, i.e. 
they are convergent, or inherited through a shared common ancester, i.e. they are synapo- 
morphic. These hypotheses are tested by comparative analysis. 

Onychostoma laticeps Gunther, 1896 (Figs 11A-C) 

The premaxilla is deeper than that of Semiplotus and has no distal ventral curvature, its 
ventral border is also thick and lacks the sharp, thin edge of the semiplotine bone. 



Fig. 11 Lower jaw bones of A, B & C, Onychostoma laticeps in lateral, dorsal and posterior 
views; D, Varicorhinus beso, dorsal view; E & F, Capoeta capoeta in medial and posterior views. 
Scale = 5 mm. 

The maxilla is deep with a broad anterior ascending process. As in Semiplotus the mid- 
lateral process is curved mesially, and although there is a posterior depression of the 
premaxilla there is no medial notch. 

The lower jaw resembles that of Semiplotus in a number of features, cf. Figs 5 & 1 1 A. The 
dentary has a bevelled labial surface, a prominent lateral ridge and a postero-ventral process. 
The coronoid process differs from the semiplotine one and that in other compared taxa in 
having a markedly concave anterior edge with a long dorso-anterior process. There is no 
synarthritic joint. The anguloarticular is broad, its posterior face shaped into a figure-8 facet. 
Although the medial portion of the bone forms an articular surface it is not posteriorly 
extended, nor does it appose a separate facet. 

The superficial jaw musculature is more complex than in Semiplotus. Adductor 
mandibulae A,b is extensive, its lower, preopercular part almost entirely tendinous; 
insertion is via a thick tendon to the lateral maxillary face. Muscle A,a is a parallel fibred 
element that inserts musculosly on the maxilla above A,b. The anterior part of the adductor 



muscle complex is crossed by a band of ligamentous tissue attached dorsally to the inner face 
of the 1st infraorbital and ventrally to the skin covering the preoperculum (Fig. 12A). Muscle 
A 2 is a narrow element, its posterior part overlying A,a (Fig. 12B). The muscle's ventral 
border is tendinous and insertion is via a thick tendon on to the posterior rim of the coronoid 
process. The area of insertion is invested by a thick connective tissue sheath which extends 
around the coronoid process and laterally attaches to the maxilla. Muscle A 3 can be 
distinguished from A 2 by its posterior separation by the levator arcus palatini muscle, and its 
separate insertion onto the coronomeckelian bone. 

MX De 

A 2 

Fig. 12 Jaw musculature of A & B, Onychostoma laticeps, A, superfcial and B, deeper muscle 
layers; C, Cyprinion microphthalmum superfcial musculature. Scale = 5 mm. 

The protractor hyoideus (Fig. 13 A) is complex and composed of several sections. The 
muscle is organised around a tendon that originates from the interoperculum and meets its 
partner at a median raphe. The anterior part of the muscle runs from the transverse tendon 
to the inner rim of the dentary. The symphysial area of the muscle grades into a thick, fatty 
connective tissue which becomes confluent with the lower lip tissue. Laterally, there is a 
separate segment of the protractor running to the hyoid arch. A dorsal portion of the muscle 
extends from the anterohyal and becomes confluent with the ventral section of the muscle. 
The posterior body of the protractor extends from the branchiostegal rays to the central, 
transverse tendon; it is dense, with its fibres, or strands of fibres widely separated by a fatty 
and tendinous tissue. The basal layer of the muscle strongly adheres to the dermis of the 
hyoid area. 



De / Phl Pop 






Fig. 13 Hyoid musculature of A, Onychostoma laticeps in oblique ventral view. Scale = 10 mm; 
B, Varicorhinus beso in oblique ventral view, and C, Capoeta capoeta direct ventral view of left 
side. B & C are semi-diagrammatic. 

The lower lip (Fig. 1 4) 

As in Semiplotus, the core of the lip is a deep, broad labial shelf of dense tissue, its dorsal 
layer (Dls) being of a more fibrous nature. Overlying, and completely free from its dorsal and 
anterior surfaces is a thick tissue envelope composed of a thin basal layer (Mlt) and a thicker, 
denser upper layer (Elt) whose lingual portion rises above the dentary rim. Anteriorly the 
epidermal layer is transformed into a keratinized sheath. Sandwiched between the labial 
shelf and the basal layer is a thin keratinized band (Mkl) that extends along the central 
portion of the labial shelf. Ventral to the shelf are layers of tissue which are the counterparts 
of those lying dorsally. The ventral dermal and epidermal layers (Vmlt, Velt) are, however, 
firmly attached to the shelf. Unlike Semiplotus, there are no tendinous bands connecting the 
dorsal lip tissue directly with the protractor hyoideus muscle and neither, apart from the 
epidermis, is the ventral tissue contiguous with that muscle. 


Mkl Elt 




Fig. 14 Sagittal sections through the lower lips of A, Onychostoma laticeps; B, Varicorhinus 
beso; C, Capoeta capoeta; D, Cyprinion macrostomum. Scale = 10 mm. 

Varicorhinus beso Ruppell, 1836 

The premaxilla is shallow and lacks a prominent anterior ascending process; the maxilla has 
broad and thick anterior, and long, mesially curved mid-lateral processes. Unlike 
Semiplotus, the posterior portion of the maxilla is not expanded, compressed or medially 

The dentary (Fig. 1 ID) has an abrupt mesial curvature with a broad labial surface and a 
laterally protruding process at the rictus; its coronoid process is tall and slopes outward. The 
anguloarticular is short and broad, its anterior tip extending beyond the base of the coronoid 
process. Unlike Semiplotus the articular facet is not developed into medial processes, and it 
is transversly aligned. 

The quadrate, like that in Semiplotus, is short and deep with an expanded articulatory 
facet. The facet is a double ellipsoid inclined at 45 to the transverse horizontal. In contrast 
to Semiplotus the facet is confined to the anterior face of the bone, no part extending to the 
lateral surface. 

The jaw musculature differs from Semiplotus in that adductor mandibulae muscle A! has 
a single tendinous insertion on the maxillary outer face. A small A w portion fans out from 
the tendon of A 3 across the anguloarticular and part of the coronoid process. The protractor 
hyoideus is strongly developed, its main portion attaching tendinously to the 1st and 2nd 



branchiostegal rays and to the posterohyal (Fig. 13B). A lateral part attaches via a strong 
tendon (almost ossiFed in large specimens) to the anterior tip of the interoperculum, a 
feature not found in Semiplotus. Its anterior part becomes tendinous and fans out to attach 
along the posterior rim of the dentary (Fig. 1 3B). 

The lower lip, although having the same basic structure as in Semiplotus has a less 
complex histological structure (Fig. 14B). The differences are a shallower labial shelf, lack of 
a basal dermal tissue layer, the dermis united at its central portion with the labial shelf, and 
more extensive keratinization of the anterior epidermis. There are no tendinous connections 
between the lip tissue and the protractor hyoideus muscle. 

Mxn -T- 

Fig. 15 

Premaxillae in dorsal view of A, Cyprinion watsoni; B, Capoeta capoeta posterior portion 

only. Scale = 5 mm. 

Capoeta capoeta Guldenstadt, 1773 

The premaxilla lacks an anterior ascending process; the maxilla has a marked mesial 
curvature, a thick and wide anterior and a shallow mid-lateral ascending process. The 
posterior arm of the maxilla, as in Semiplotus, is compressed and medially notched (Fig 1 5). 

The dentary has the same abrupt medial curvature and antero-ventrally sloped labial 
surface as in Varicorhinus beso (Fig. HE). Unlike Semiplotus, Onychostoma and 
Varicorhinus, the coronoid process is aligned at 45 to the midline, so that it is set at the 
curve of the dentary. The ventro-posterior surface of the dentary is broad, as is the depressed 
anguloarticular which overlies it. The articulatory facet is ovate and transversly aligned (Fig. 

The jaw musculature of Capoeta is similar to that of Semiplotus and Varicorhinus. 
Adductor mandibulae A, is a single muscle with a central anterior aponeurosis; insertion is 
via a thick tendon onto a lateral maxillary process. A thin tendinous strip of tissue runs from 
the dorso-anterior part of the muscle to insert on the connective tissue covering the medial 
face of the maxilla. Muscle A 2 inserts entirely on the posterior border of the dentary coronoid 
process; a separate group of fibres leaves the main tendon to run ventrally across the medial 
coronoid face. Tendon A 3 inserts on the coronomeckelian bone. 

The protractor hyoideus is extensively tendinous in its posterior part and it is attached to 
all the hyoid elements except the interhyal. There is no lateral portion stemming from the 
interoperculum as in Varicorhinus. Anteriorly the left and right arms of the muscle are 
widely separated so that the intermandibularis is exposed (Fig. 1 3C). 

The lower lip differs from that in the other taxa compared here in having a narrow labial 
shelf, no layered organisation of the tissue overlying the shelf, and this tissue continuous with 



that lining the mouth. The outer, keratinized sheath lies along the upper edge of the labial 
shelf rather than around its anterior face. 

Cyprinion Heckel, 1843 

In Cyprinion species, the premaxilla usually lacks an ascending process, and when one is 
present it is but slightly developed. Maxillary shape is interspecifically variable and in only 
one species, C. watsoni, does it closely resemble Semiplotus in having a reduced, medially 
directed mid-lateral ascending process and a depressed, medially notched posterior arm (Fig. 

The lower jaw of C. macrostomum and C. microphthalmum resembles that of Semiplotus 
more closely than does that of any other Cyprinion species. This resemblance is in the 


Fig. 16 Lower jaw bones (right side) of A & B, Cyprinion acinaces in dorsal and medial views; C, 
Cyprinion watsoni in dorsal view; D & E, Cyprinion microphthalmum in dorsal and posterior 
views; F, quadrate in lateral view; G & H, Cyprinion macrostomum in dorsal and posterior 
views; I, quadrate in anterior view; J & K Cyprinion kais in dorsal (right jaw bone) and lateral 
(left jaw bone) views. Scale = 5 mm. 

320 G. J. HOWES 

curvature of the dentary, a complex synarthritic joint and the broad articular surface of the 
anguloarticular. There is a slight medial process, equivalent to process Apl in Semiplotus, 
separated from the lateral part of the bone by a deep cleft. Both Cyprinion species also 
possess a quadrate bearing part of its articulatory facet laterally (Fig. 16F). Cyprinion watsoni 
has a more elongate jaw than C. microphthalmum and C. macrostomum, but the angulo- 
articular has a similar broadening of its facet. 

With the exception of C. microphthalmum the muscles attaching to the upper jaw are 
undivided or only partially divided. In that species the adductor mandibulae A,a is separated 
from A,b for almost its entire length but shares a common insertion with it on both the outer 
and inner aspects of the maxilla (Fig. 12C). The lower jaw muscles are also interspecifically 
variable with respect to their proportions and fibre orientation, but their insertions are 
essentially similar. There is some variability in the size of muscle A w . In C. microphthalmum 
and C. macrostoma, as in Semiplotus it inserts on a medial tendon of muscle A 2 . 

The lower jaws of C. acinaces, C. milesi and C. watsoni are less derived with respect to 
their labial and articulatory surfaces but all have a synarthritic joint of varying complexity, 
the weakest being in C. acinaces (Fig. 16A). The jaw of C. acinaces is regarded as the most 
plesiomorph of all Cyprinion species with respect to its moderately expanded labial surface, 
mid-laterally placed coronoid process and small, transverse articulatory facet. The lower jaw 
of C. kais is deeper posteriorly than in any other Cyprinion species, and its labial surface is 
much narrower (Fig. 16J); as in Semiplotus, the anguloarticular covers a substantial area of 
the medial face of the dentary coronoid process. In all Cyprinion species, the anguloarticular 
extends posteriorly only marginally beyond the dentary. 

The lower lip of Cyprinion is of the same layered type found in Semiplotus (Fig. 14D), 
with that of C. macrostoma having the greatest resemblance in terms of dimensions of the 
labial shelf, tissue layering, and tendinous connections with the protractor hyoideus muscles. 
The degree of keratinization is interspecifically variable but reaches its greatest development 
in C. macrostomum. 

Onychostoma, Varicorhinus and Capoeta all lack the Semiplotus features of a synarthritic 
mandibular joint, medial anguloarticular processes, lateral quadrate facet, aponeurotic 
connection of the adductor mandibulae and protractor hyoideus muscles, interhyal 
connection of the protractor hyoideus, and complexity of lower lip structure. Onychostoma 
shares with Semiplotus a lateral dentary ridge and posteroventral process, but the nature of 
the coronoid process and the lack of the apomorphies cited above favour the hypothesis that 
the derived jaw characters held in common have been independently acquired. Likewise, the 
sector type of mouth morphology of Varicorhinus and Capoeta is believed to be a convergent 

The phyletic relationships of Onychostoma, Varicorhinus and Capoeta have yet to be 
determined, but other apomorphies (absent in Semiplotus and Cyprinion) suggest the 
following affinities: Onychostoma is related to Ageniogarra and possibly to a group of 
Chinese 'Schizothorax*; Varicorhinus forms a close relationship with certain African Barbus 
(Banister, in preparation); Capoeta is the likely sister-group to Cyprinus and Carassius (see 
Howes, 1981). 

As noted above all Cyprinion species possess a synarthritic mandibular joint, aponeurotic 
connection of the adductor mandibulae and protractor hyoideus muscles and similarly 
developed lower lip structure. Synapomorphies other than those of the jaws also suggest a 
close relationship between Semiplotus and Cyprinion. 

Other apomorph characters of Semiplotus 

In addition to the jaws, Semiplotus species are characterized by other and presumed 
apomorphic features. 

1 . Condylar articulation of the interhyal with the symplectic. The usual condition in 
cyprinids is for the symplectic to be an elongate bone. It is rarely short and deep, and 



only in a minority of taxa is the posterior face concave and in articulation with the 
interhyal. The common condition of the interhyal is a lamellate bone, sometimes 
incompletely ossified. 

In Semiplotus the symplectic is short and buttress-like (Figs 6 & 1 7A), its posterior 
surface is concave and articulates directly with the interhyal, thereby forming part of 
the ball and socket joint between the two elements. The interhyal is short and 
cylindrical, and articulates with the dorsal surface of the posterohyal via a cartilaginous 


Fig. 17 Hyoid and suspensorial elements of A & B, Semiplotus semiplotus, medial and 
dorsal views of posterior hyoid bar; C, Cyprinion macrostomum, medial view of posterior hyoid 
bar; D & E, Varicorhinus beso, medial and dorsal views of posterior hyoid bar; F-I, symplectic 
bones of F, Cyprinion microphthlamum; G, C. kais; H, C. acinaces; I, C. watsoni; J, Cyprinion 
kais, anterior view of 1st branchiostegal ray and hyoid bar. Scale = 3 mm. 

322 G. J. HOWES 

Amongst the taxa considered here, only in Onychostoma and some species of 
Cyprinion does the symplectic approach that of Semiplotus in size and shape. In 
Onychostoma, the condylar surface of the bone faces ventrally whereas in Semiplotus 
and Cyprinion it is angled postero- ventrally. The interhyal in all these genera, and in 
Varicorhinus, is hypertrophied and articulates with the dorsal aspect of the postero- 
hyal, but since there are no other synapomorphies of the hyoid elements shared by 
Onychostoma, Varicorhinus and Semiplotus, these are considered to be independently 
derived features (see p. 320). 

2. Ventral and lateral articulation of the 3rd branchiostegal ray with the posterohyal (Figs 
17A & B). Plesiomorphically the 3rd branchiostegal is attached ligamentously to the 
lateral face of the posterohyal. In Semiplotus, the posterohyal is short and thick, and 
the branchiostegal ray expanded medially so as to form a shelf underlying the ventral 
border of the posterohyal with whose posterior portion it articulates. The anterior part 
of the branchiostegal shelf is attached ligamentously to the medial face of the postero- 
hyal whilst the proximal lamellate area of the branchiostegal ray is attached to the 
lateral face of the hyal. 

A slight medial shelf is present on the 3rd branchiostegal rays of Varicorhinus and 
Capoeta. It is, however, barely developed in these taxa and the posterohyal is not 
distally expanded as in Semiplotus (Fig. 17). Such an expansion of the branchiostegal is 
lacking in Onychostoma. Only in Cyprinion is there a substantial 3rd branchiostegal 
shelf, which in C. macrostomum is developed much as in Semiplotus (Fig. 1 7). 

3. Ventrally directed supraethmoid and hypertrophied vomer (Figs 1 8A-C). Although the 
rostral curvature of the ethmoid bloc is variable in the Cyprinidae (see Howes, 1980) 
nowhere is it more extreme than in Semiplotus where the supraethmoid is almost 
vertically aligned. An apparent correlate of this condition is the hypertrophied vomer, 
whose enlargement also results in a steep angle forming between it and the para- 
sphenoid (Fig. ISA). 

In both Onychostoma and Varicorhinus the ethmoid is strongly curved ventrally and 
there is a corresponding hypertrophy of the vomer. In these genera the ventrally 
directed part of the supraethmoid is confined to the lower part of the bone. In 
Cyprinion macrostomum the curvature of the supraethmoid more closely resembles the 
condition in Semiplotus where the entire bone slopes ventrally. 

4. Frontal- supraethmoid foramen (Figs 18B-C). Synapomorphic for Semiplotus semi- 
plotus and S. burmanicus is a foramen between the frontals and the supraethmoid (see 
p. 304). The posterior border of the foramen indents each frontal to form a notch in 
much the same way as the antero-medial supraethmoid notch occurs in other cyprinids. 
Likewise, the frontal notch of Semiplotus performs a similar function, namely in 
providing a hold-fast for the kinethmoid ligaments. Attachment of kinethmoid 
ligaments to the frontal instead of the supraethmoid is unknown in any other cyprinid. 

5. Elliptical subtemporal fossa with major contribution from the pterotic (Fig. 18D). The 
plesiomorph subtemporal fossa is shallow with a round or oval outline (see Howes, 
1981). In Semiplotus the fossa is deep and markedly ellipsoidal, its long axis transverse. 
Only in Onychostoma does the subtemporal fossa approach the shape of that in S. 
semiplotus and S. burmanicus, but it lacks the extreme depth of those species. 

Typically the posterior wall of the subtemporal fossa in cyprinids is formed princi- 
pally by the exoccipital, with a minor contribution from the pterotic (see, for example, 
Howes, 1978, Figs 6 & 33; 1980, Fig. 13). In Semiplotus the greater part of the posterior 
wall is formed by the pterotic (Fig. 18D); only in Cyprinion is there a similarly 
substantial contribution of the pterotic. 

6. Proximal part ofsupraoccipital crest formed by the parietals (Fig. 1 9 A). 
Semiplotus species have a high supraoccipital crest; the parietals are short and slope 
upwards, their medial portions extending posteriorly and embracing the base of the 
supraocipital crest. With the exception of Cyprinion, in no other cyprinid taxon, even 





Fig. 18 Semiplotus semiplotus, cranium. A, lateral view of ethmo-vomerine and orbital areas; B, 
dorsal & C, anterior views of ethmoid region; D, ventral view of posterior cranial area; E & F, 
basioccipital process in lateral view of E, S. semiplotus and F, S. burmanicus. Scale = 10 mm. 

those with a relatively high supraoccipital crest, does the parietal extend postero- 
medially. In Cyprinion there is some variability in the length of the parietal, but in all 
species it extends postero-medially to form part of the supraoccipital crest (Figs 

Reduced basioccipital process and masticatory plate (Figs 1 8E & F). 
Although a reduction of the basioccipital process and masticatory plate occurs in 
several groups of cyprinids (see Howes, 1981), in none is the reduction so extreme as in 
Semiplotus. In S. burmanicus, the basioccipital (masticatory) plate is entirely absent 
and the ventral (aortic) ossification is transversly convex. Only in some Cyprinion 
species (C. kais and C. microphthalmum) is a truncated basioccipital process associated 
with marked reduction of the ventral plate. 
Interoperculum-preoperculum articulation (Figs. 6B-C). 

In Semiplotus the anterior tip of the preoperculum is formed into a condyle and is 
curved medially so as to overlie the anterodorsal part of the interoperculum; the 





Fig. 19 Dorsal posterior cranium of A, Semiplotus semiplotus; B, Cyprinion macrostomum; C, 

Cyprinion kais. Scale = 5 mm. 

anterior tip of the interoperculum is formed into a facet. The two elements are firmly 
articulated and the medial curvature of the preoperculum beneath the quadrate appears 
to reinforce the rigidity of the quadrate-lower jaw articulation. The interoperculum 
also bears a medial facet with which the rounded tip of the posterohyal articulates. 

In Varicorhinus there is a similar development of the terminal point of the intero- 
perculum, but only in Cyprinion do both elements articulate as in Semiplotus (Fig. 
161). A medial interopercular facet is variously developed in the Cyprinidae, but in 
Cyprinion alone is it of similar depth and position. 

Development of the supraneurals and articulation with 1st dorsal pterygiophore (Fig. 20). 
The supraneurals of Semiplotus are hypertrophied and each is in contact with its 
respective neural spine. The 'neural complex' of the Weberian apparatus is upright and 
has an almost straight anterior edge; dorsally it contacts the posteriorly directed supra- 
occipital process. This is contrary to the usual cyprinid condition where the anterior 
border of the neural complex is concave, giving it an axe-shaped appearance. In 
Semiplotus semiplotus there are 5 or 6 supraneurals, each articulating with its neigh- 
bour; the posterior element is bifurcated and overlaps the anterior margin of the 1 st 
dorsal pterygiophore. Semiplotus burmanicus has 7 supraneurals, all somewhat thinner 
than in S. semiplotus but similarly and sequentially joined, each bearing a vertical ridge 
laterally. The 1st supraneural is narrowly separated from the neural complex. 

Although enlarged supraneurals occur in other cyprinid taxa they are never so tall as 
those in Semiplotus semiplotus. Apart from S. burmanicus, interlocking of the last 
supraneural with the 1st pterygiophore occurs only in Cyprinion macrostomum, and a 
straight-edged neural complex is shared only with Cyprinion microphthalmum. 






Fig. 20 Anterior vertebral column of A, Semiplotus semiplotus; B, S. burmanicus. Scale = 5 mm. 

The relationship between Semiplotus and Cyprinion 

In addition to the jaw synapomorphies (p. 320), Semiplotus shares with Cyprinion species a 
series of derived hyoid, cranial and vertebral features (enumerated above as 1-9), some of 
which are present in all Cyprinion (I, 2, 5, 6, 8), others in only some species of that genus 

At present, the genus Cyprinion cannot be defined on the basis of even a single autapo- 
morphy. Derived characters such as the expansion of the proximal portion of the pelvic fin 
rays, interpelvic papillate flaps (see Banister & Clarke, 1 977) and a naked pre-dorsal ridge 
(see Mirza, 1969) although synapomorphic for some species do not occur in all members of 
the genus. If, however, the concept of Cyprinion is expanded so as to include Semiplotus, 
then the taxon so formed is recognisable as a monophyletic unit on the basis of several 
synapomorphies (the principal one being the presence of a synarthritic dentary joint), and 
the problem posed by those apomorphies shared by Semiplotus and some Cyprinion species 
is also overcome. Thus (following Heckel, 1843; see p. 331), Semiplotus is regarded as a 
junior synonym of Cyprinion and any references to Cyprinion from here onwards must be 
taken to include Semiplotus semiplotus and S. burmanicus. (A synopsis of included taxa is 
given on p. 331.) 

326 G. J. HOWES 


Apomorph character distribution in Cyprinion species 

If the jaws of Cyprinion semiplotum are seen as the terminal stage of increasing complexity, 
then there is a possibility that an evolutionary or 'transitional sequence' of jaw types might 
reflect the pattern of interspecific relationships within the genus Cyprinion. However, such 
an arrangement of increasingly derived jaw morphology (in terms of hypertrophy or 
reduction of various elements) is not congruent with polarized morphoclines in other 
characters amongst Cyprinion species. 

Jaw synapomorphies possessed by C. semiplotum and C. burmanica unite them with other 
Cyprinion species in the following patterns: 

(i) A greatly increased complexity of the mandibular synarthritic joint, a broad angulo- 
articular with a medial process and a cleft between the lateral and medial faces of the 
bone. Shared with C. macrostomum, C. microphthalmum and C. watsoni. 
(ii) A lateral quadrate facet and a medial A 2 tendon. Shared with C. macrostomum and C. 


(iii) A medial maxillary notch. Shared with C. watsoni. 

(iv) A divided insertion of adductor mandibulae A, muscle. Shared with C. micro- 

Any 'transitional series' present amongst this character complex is interrupted by the 
distribution of characters (iii) and (iv). The following synapomorphies uniting C. semi- 
plotum and C. burmanica among other Cyprinion species display incongruent patterns 
when set against those of the jaw: 

(v) A deep symplectic and a truncated basioccipital process with a reduced masticatory 
plate. Shared with C. kais and C. microphthalmum. 

(vi) A straight-edged neural complex. Shared with C. microphthalmum. 

(vii) Articulation of the last supraneural with the 1 st pterygiophore. 
Shared with C. macrostomum. 

Synapomorphies which are not present in C. semiplotum and C. burmanica unite 
the following species: 

(viii) 1st branchiostegal ray articulating ventrally with the ceratohyal through a 
long medial process (Fig. 17). Shared only by C. macrostomum and C. kais. 
(Cyprinion kais, it should be remembered has neither of the derived jaw 
characters (i) nor (ii) which C. macrostomum shares with other species). 

(ix) Pelvic flaps and papillate interpelvic region. Shared by C. watsoni, C. acinaces and 
C. milesi. The two latter species are conservative in their cranial and jaw 
morphologies (see p. 320) and are thought to comprise the plesiomorph lineage 
within the genus. Thus, unlike C. watsoni, C. acinaces and C. milesi share none of 
those characters that distinguish the more derived lineage (see (i) and (ii) above). 

In virtually all monophyletic assemblages of cyprinid fishes so far identified there are sets 
of incongruent apomorphies, a state of affairs which makes it impossible to apply a rigidly 
dichotomous system of relationships. Such incongruencies usually can be ascribed to some 
form of homoplasy, particularly when the characters in question occur in outgroups, as for 
example the medial maxillary notch present in Cyprinion semiplotum, C. burmanica, C. 
watsoni and also in Capoeta. Here the most likely explanation for its presence in Capoeta 
and Cyprinion is one of parallelism as no other synapomorphies unite Capoeta with 
Cyprinion; see p. 320. However, its distribution within the three Cyprinion species can only 
be accounted for most parsimoniously by assuming its loss in two other species (C. micro- 
phthalmum and C. macrostomum). 

Other character sets also reveal incongruency when treated dichotomously. For example, 
in Cyprinion kais one must postulate the loss of all jaw synapomorphies plus the parallel 
evolution in this species and in C. semiplotum, C. burmanica and C. microphthlamum of 
similarly derived symplectic and basioccipital characters. Furthermore, the loss of a 



supraneural-pterygiophore articulation must be assumed in C. kais and C. micro- 
phthalmum, and interpelvic papillate flaps must be treated either as a plesiomorph character 
or assumed to be secondarily derived in C. watsoni. Such solutions result in a confusing 
mixture of assumed parallelism and reversal, with the acceptance of one alternative being no 
more parsimonious than the acceptance of another; see cladogram, Fig. 2 1 . 

Bremer & Wanntorp (1979) have discussed the difficulties posed by conflicting synapo- 
morphies and show that the situation may best be displayed as reticulate cladograms. Such a 
device is used here to show character distribution in Cyprinion (Fig. 22). 

Although sister species cannot be identified with this synapomorphy scheme, both 
apomorph character weighting and the simplistic, essentially numerical approach of using 

Fig. 21 Most parsimonious cladogram of Cyprinion characters. Synapomorphies; nodes 1, 
Cyprinion synapomorphies (including expanded pelvic fin rays); 2, jaw synapomorphies (see 
text, p. 320); 3, lateral quadrate facet, medial A 2 tendon, truncated basioccipital, deep 
symplectic; 4, upright neural complex; 5, jaw and cranial synapomorphies (see text); 6, inter- 
pelvic papillate flaps; 7, method of articulation of 1st branchiostegal with ceratohyal (see text, 
p. 326). Parallelisms and reversals; lineage A, scaleless interpelvic region (parallel to 6); B, loss 
of deep symplectic and truncated basioccipital in C. macrostomum, parallelism of 
pterygiophore-supraneural articulation to C. semiplotus and C. burmanicus; C, loss of jaw 
synapomorphies in C. kais; D, loss of supraneural-pterygiophore articulation in C. 
microphthalmum; E, loss of pelvic fin ray expansion in C. semiplotus and C. burmanicus. 



Fig. 22 Reticulate synapomorphy scheme for Cyprinion characters Level 1, Cyprinion 
synapomorphies (see text, p. 325); 2, expanded pelvic fin rays; 3, increased complexity of 
synarthritic joint, broadened anguloarticular with medial process Ap 3 ; 4, lateral quadrate facet, 
medial A 2 tendon, interopercular-preopercular articulation; 5, jaw and cranial synapomorphies 
(see text, p. 312). Conflicting synapomorphies: (a) interpelvic papillate flaps; (b) articulation of 
1st branchiostegal ray; (c) upright neural complex; (d) divided adductor mandibulae A,; (e) 
medial maxillary notch; (h) supraneural-pterygiophore articulation. Black boxes indicate the 
presence of that character on a particular lineage, a break in the horizontal indicates its absence. 


the greatest number of synapomorphies as an assessment of relationship is avoided 
(Tarsitano & Hecht, 1980: 178). Likewise, the alternative approach, using unresolved 
polychotomies, serves only to indicate that there is an absence of characters rather than the 
presence of conflicting ones. 

It may be argued that the choice of character sets employed here is itself a weighting 
procedure, and that the use of other sets would reveal dichotomous sister-group relation- 
ships. The counterclaim to these arguments is that the characters employed are those whose 
intraspecific variability is known, and indeed are the only definitive characters recognizable 
at this level of investigation. Cyprinids are 'notorious' for their conservative characters (see 
for example Regan, 1911). Banister (1980) held that this conservatism '. . . coupled with the 
occasional small saltatory morphological changes . . . has the effect of making it difficult, if 
not impossible to distinguish between parallelism and the sharing of characters derived 
through common ancestry'. In each of the cyprinid groups so far identified as monphyletic 
(the aspinines, chelines, bariliines and abramines) one lineage appears as the bearer of a large 
number of autapomorphies and thus leaves a considerable 'morphological gap' between it 
and the more conservative corpus of lineages comprising that particular group. Examples are 
Macrochirichthys + other chelines and Hypophthalmichthys + other abramines (see Howes, 
1979; 1981). Such also appears to be the case within Cyprinion where C. semiplotum and C. 
burmanica represent a highly derived lineage characterized by many jaw synapomorphies. 
Presumably the accumulation of synapomorphies in one such lineage is the result of a 
cumulative series of speciation events, lacking in the other, more generalized lineages (i.e. 
'the occasional small saltatory changes' of Banister, 1980). 

Whether the reticulate pattern of relationships is an artifact resulting from incomplete 
character analysis or a reflection of polymorphic descent through populations of the intra- 
group lineages is a problem as yet unresolved. 

Evolution of the Cyprinion jaw and the 'sector mouth' 

Even though no 'transitional series' of jaw types emerges from the reticulate pattern 
presented (Fig. 22), particular synapomorphies indicate a possible evolutionary course 
leading to the specialized morphology of the Cyprinion semiplotum-C. burmanica lineage. 
Comparison of these apomorphies with similar but independently derived characters in 
other cyprinid lineages indicates those features which are prerequisites for the development 
of a sector mouth. 

In the upper jaw a decisive step towards a sector mouth appears to have been taken with 
the reduction of the premaxillary ascending process. Matthes (1963) remarks that 
reduction of this process probably occurred independently in several genera and is not there- 
fore indicative of close relationship. Matthes was referring particularly to similarities 
between Varicorhinus, Labeo and Garra. Whilst this study endorses Matthes' view that this 
particular feature is not necessarily a synapomorphy for all three genera, it does not refute its 
possible synapomorphic status in Labeo and Garra, which share other derived characters 
(Reid, unpublished thesis). 

At first sight it would seem that reduction of the premaxillary processes should lead to 
restricted protrusibility of the upper jaw. However, this may be compensated for by a 
pronounced downward curvature of the ethmoid bloc as seen in Cyprinion and 
Onychostoma and, to a lesser extent, in Varicorhinus and Capoeta. The depressed ethmoid 
region in Labeo and Garra possibly fulfills the same function. 

The reduced and medially curved mid-lateral ascending maxillary process of Cyprinion 
semiplotum appears otherwise only in Onychostoma. In both taxa this feature is apparently 
correlated with an extensive area of connective and tendinous tissue attaching to the inner 
face of the 1st infraorbital (see p. 305). The medial notch on the posterior arm of the maxilla 
in some Cyprinion species and in Capoeta performs a similar function as a hold-fast for the 
thick tissue connecting the maxilla with the coronoid process of the dentary (see p. 305). 

Concerning the Cyprinion lower jaw, the most important evolutionary innovation appears 

330 G. J. HOWES 

to have been the development of a synarthritic mandibular joint. Such a joint usually 
suggests some degree of lateral movement, as in the 'knuckle joint' of cynodontine 
characoids (Howes, 1976), but in the case of Cyprinion it may act as a compensatory device 
to the torque induced by the contraction of the thickly ligamentous protractor hyoideus 
muscle. The absence of any lateral jaw movement is also suggested by the strongly produced 
lateral and medial anguloarticular processes which provide a rigid coupling with the 

Changes associated with increased medial curvature of the dentary appear to be manifest 
in the outward slope of the coronoid process so that the axis and insertion angle of the 
adductor mandibulae A 2 and A 3 muscles remain unchanged. This is contrary to the 
condition in Capoeta where the coronoid process and muscle fibres are set at an angle to the 
body axis. In Cyprinion there is a noticeable change in the relative position of the coronoid 
process from its mid-lateral point in the plesiomorph C. acinaces to a posterior position in C. 
semiplotum. This apparent positional shift is due to morphological changes in the 
surrounding elements, i.e. broadening and posterior shortening of the anguloarticular and 
the medial curvature of the dentary (Fig. 1 6 A-E, G-K). 

The rigid articulation of the lower jaw in Cyprinion semiplotum and C. burmanica 
appears to restrict its vertical movement and it seems merely to act as a firm platform for the 
highly mobile lip. 

The lower lip of Cyprinion differs from that in other taxa in being a mobile structure; the 
epidermis is so loosely attached to the underlying labial shelf that it is free to move antero- 
posteriorly, and its mobility appears to be controlled by the action of the protractor hyoideus 
complex via dorsal and ventral tendinous linkages. This additional function of the protractor 
hyoideus is also indicated by a multi-directional joint between the hyoid and the 

Thus, in the more derived Cyprinion species, increased complexity of the jaw-quadrate 
articulatory surfaces and of the lower lip is correlated with increased mobility of the hyoid 
bar and hypertrophy of the protractor hyoideus. 

Where it occurs, a sector mouth appears to have the same basic function i.e. for scraping or 
ploughing epilithic material (see Matthes, 1963). Certainly, there are some 'necessary design 
components' common to all lineages of taxa with a sector mouth, viz. marked medial 
curvature of the anterior portion of the dentary; a broadened anguloarticular and barely 
extending beyond the posterior border of the dentary; greater mobility of the hyoid- 
suspensorial joint; 3rd branchiostegal ray shifting ventrally to the posterohyal, and an hyper- 
trophy of the protractor hyoideus musculature. 

Apart from Cyprinion such specializations can be found among Barbus-Varicorhinus 
species. Doubtless, cranial and vertebral apomorphies (deepening of the posterior cranial 
region and increased rigidity of the anterior vertebral column) form a functional unity 
contributing to epilithic feeding activity. Just how these components interact functionally 
with those of the jaws, the hyoid and branchial arches, are problems to be solved. 

An earlier remark (Howes, 1981) that the '. . . morphology of the lower jaw is possibly the 
most variable unit in Cyprinidae . . .' is borne out by this study. That the cyprinid jaw 
appears to be so adaptable suggests that, as in cichlids, the pharyngeal bones perform a 
dominant functional role of food manipulation and mastication, so freeing the jaws for 
specialized food gathering. No comparative experimental functional data are available for 
cyprinid taxa and it is not known whether the pharyngeal apparatus is as versatile as that of 
cichlids and whether it exerts a primary dominant influence on jaw modification. Liem 
(1980) has pointed out that in cichlids epilithic feeding patterns are of a kind requiring 
specializations that exceed those of simple inertial suction feeding. Certainly this appears to 
be true for sector-mouthed cyprinids as well, where (in the more derived forms) upper jaw 
protrusion is minimal and the manipulative function of the jaw in food gathering has been 
transferred, in part, to the hyoid arch. 


Synopsis of Cyprinion species 

The synonymies include only the original citation of the species. 
CYPRINION deckel, 1843 

Semiplotus Bleeker, 1863 
Scaphiodonichthys Vinciguerra, 1889 
Scaphiodontopsis Fowler, 1934 

The reasons for synonymising Semiplotus with Cyprinion have been explained in this paper 
p. 299). 

Scaphiodonichthys was separated from 'Semiplotus' on the differences in the number of 
branched dorsal fin rays (9-12 in Scaphiodonichthys, 20-25 in 'Semiplotus'). Smith 
(1945 : 206) quotes from a letter he received from Dr S. L. Hora who suggested that inter- 
mediate forms were '. . . yet to be discovered with a number of branched rays intermediate 
between 12 and 20. When such forms are found Semiplotus and Scaphiodonichthys will have 
to be regarded as congeneric . . .' To my knowledge no such 'intermediate' forms have been 
found. The existence of synapomorphic osteological and myological characters in species of 
both 'Scaphiodonichthys'' and Cyprinion indicates their generic unity (see p. 312 for a list of 
characters). When viewed as species of Cyprinion, then 'intermediate forms' linking 
''Semiplotus'' and 'Scaphiodonichthys' are in fact present, and indeed the number of branched 
dorsal fin rays does range from 10-25. 

Species occurring east of the Himalayas 

Cyprinion semiplotum McClelland, 1839 

Cyprinus semiplotus McClelland, 1839 Asiat. Reschs. 19 : 374 
Cyprinion semiplotus Heckel, 1843 in Russegger's Reisen 1 : 1015 
Semiplotus semiplotus Bleeker, 1 863 Atlas Ichth. 3 : 25 
Semiplotus mcclellandi Bleeker, 1863 Atlas Ichth. 3 : 25 
Semiplotus cirrhosusChaudhuri, 1919 Rec. Ind. Mus. 16 : 280 

DISTRIBUTION. Nepal, north Bengal (see Hora, 1937; Hora & Gupta, 1940). 

Cyprinion modestum Day, 1870 

Semiplotus modestus Day, 1870 Proc. Zool. Soc. Lond. : 101 

Day (1870) considered this species as '. . . intermediate between the genera Semiplotus and 
Cyprinion'. Unlike C. semiplotum but in common with other Cyprinion species, C. 
modestum has a serrated dorsal fin spine. No specimens have been examined in the course of 
this study (the presumed types are in Calcutta) but according to Hora (1937 : 46) Day's 
species is 'quite distinct'. 

DISTRIBUTION. Akyab Hills, Upper Burma. 

Cyprinion burmanica Vinciguerra, 1889 

Scaphiodonichthys burmanicus Vinciguerra, 1 889 Ann. Mus. Civ. Storia Nat. Geneva (2) 9 : 285 
Scaphiodontopsis acanthopertus Fowler, 1934 Proc. Acad. Nat. Sci. Philad. 86 : 119 
Onychostoma macroacanthus Pellegrin & Chevey, 1936 Bull. Soc. Zool. France 6\ (1) : 18-27 

Taki (1975) demonstrated an overlap of lateral line scale numbers between C. burmanicus 
and C. acanthopterum and stated that the two species were distinguishable only by the 
number of dorsal fin rays. I find that in a sample of 16 specimens identified as C. burmanica 
(28-65 mm SL uncat. Smithsonian Coll. Upper Thailand), there are nine specimens with 1 1 
and seven with 10 branched dorsal rays. There are thus no meristic differences separating the 
two 'species'. Since I can find no morphological differences between the syntypes of C. 
burmanica and the series of C. acanthopterum studied by Smith (1945), I consider the 
species to be synonymous. 

332 G. J. HOWES 

DISTRIBUTION. Burma, Thailand, Laos, Vietnam. 

Species west of the Himalayas 

Despite the revision of Berg (1949) and the work of Mirza (1969), many taxonomic problems 
remain concerning the species in this area. As yet, no author has examined all the nominal 
types. Also, too few specimens have been examined to judge the degree of intraspecific 
variability in certain characters. 

Cyprinion acinaces Banister & Clarke, 1 977 

Cyprinion acinaces Banister & Clarke, 1977 J. Oman Studies : 123-126 


Cyprinion macrostomum Heckel, 1843; type species of the genus 

Cyprinion macrostomum Heckel, 1843 Ichthyologiein Russegger's Reisen 1 : 1065 
?Cyprinion neglectum Heckel, 1 846 Die Fische Persiens in Russegger's Reisen 2 (3) : 223 
?Cyprinion tenuiradius Heckel, 1 846 Die Fische Persiens in Russegger's Reisen 2 (3) : 26 1 

Berg (1949) included C. neglectum in synonymy without having seen the type and regarded 
C. tenuiradius as a valid species. However, from Heckel 's figure C. tenuiradius would seem 
to be simply a 'variant' of C. macrostomum. 

DISTRIBUTION. Tigris-Euphrates (see Banister, 1980). 

Cyprinion kais Heckel, 1 843 

Cyprinion kais Heckel, 1843 Ichthyologiein Russegger's Reisen 1 : 1066 
?Cyprinion cypris Heckel, 1843 Ichthyologie in Russegger's Reisen 1 : 1067 

Berg (1949) included C. kais in the synonymy of C. macrostomum on the grounds of minor 
variation in position of the dorsal fin with respect to the pelvic fin insertion (the character 
Heckel had used to separate the species). Although Berg (1949) recognised differences in 
mouth shape between the taxa, he obviously considered them to be of little taxonomic worth. 
I have seen the types of neither C. macrostomum nor C. kais, and the recognition of these 
species is based on a comparison of Heckel's (1843) figures with two recognisably distinct 
taxa represented in the BMNH collections. In C. kais the mouth is small, with the 
keratinized lip of the lower jaw prominent and more upwardly directed than in C. macro- 
stomum. Other differences involving the morphology of the jaws and hyoid elements are 
given elsewhere in this paper. 

From Heckel's (1 843) figure (Fig. 3, pi. 7) it seems likely that C. cypris is synonymous with 
C. kais rather than with C. macrostomum as indicated by Berg (1949), and that it represents 
a juvenile specimen in which the keratinization of the jaw is still incomplete. 

DISTRIBUTION. Tigris-Euphrates. 

Cyprinion microphthalmum Day, 1880 

Scaphiodon microphthalmus Day, 1880 Proc. Zool. Soc. Lond. : 227 
?Scaphiodon muscatensis Boulenger, 1887 Proc. Zool. Soc. Lond. : 665 
Cirrhina afghana Gunther, 1889 Trans. Linn. Soc. Zool. 5 : 106 
Cirrhina afghana nikolskii Berg, 1905 Ann. Mus. Zool. Acad. Sci. 10 : 106 
Scaphiodon baluchiorum Jenkins, 1910 Rec. Ind. Mus. 5 : 124 

This synonymy is that of Berg (1949) but appears suspect. The type specimens of 
Scaphiodon microphthalmus are possibly lost (see Banister & Clarke, 1977) and until such 
time as they reappear certain identification of this species is not possible. However, Mirza's 
(1969) description based on specimens from Quetta, the type locality, appears to represent 
the species Day (1880) described. Specimens I have examined from Baluchistan (BMNH 


1883.8.2 : 20-26) and the syntypes of Cirrhina afghana (BMNH 1886.9.12 : 155-159; 21) 
conform to Mirza's description and that given by Jenkins (1910) for Scaphiodon 

Banister & Clarke (1977) followed Berg (1949) in regarding Cyprinion muscatensis as 
synonymous with C. microphthalmum. 
DISTRIBUTION. Baluchistan, Afghanistan, the Indus Plain and Saudi Arabia. 

Cyprinion milesi Day, 1 880 

Barbus milesi Day, 1 880 Proc. Zool. Soc. Lond : 228 
Barbus bampurensis Nikolsky (1899) 1900 Ann. Mus. St. Petersb. 4:410 
Scaphiodon daukesi Zugmayer, \9\2Ann. Mag. nat. Hist. 8 : 596 
Barbus baschakirdi Holly, 1929 Anz. Akad. Wiss Wienl : 1 
Cyprinion milesi Berg, 1949 Trudy zool. Inst. Leningr. 8 (4) : 821 

DISTRIBUTION. West Pakistan and Iran (see Mirza, 1969): BMNH specimens from 
Afghanistan (1889.2.1 : 263-4) and Dizak, Baluchistan (1883.8.2 : 2-3). 

Cyprinion watsoni Day, 1872 

Scaphiodon watsoni Day, 1872 J. Asiatic Soc. Bengal 41 : 324 

Scaphiodon irregularis Day, 1872 J. Asiatic Soc. Bengal 4\ : 321 

ICyprinion kirmanse Nikolski, 1899 Ann. Mus. St. Petersb. 4:412 

Scaphiodon macmahoni Regan, 1906 J. Asiatic Soc. Bengali : 8 

Scaphiodon watsoni var. belensis Zugmayer, \9\2Ann. Mag. nat. Hist. (8) 10 : 596 

Scaphiodon readingi Hora, 1923 Rec. Ind. Mus. 25 : 379-382 

Cyprinion watsoni Berg, 1949 Trudy zool. Inst. Leningr. 8 (4) : 814 

The above synonymy mainly follows that of Mirza (1969) who found a complex overlap of 
morphometric and meristic characters between C. watsoni and Scaphiodon irregularis. 
Certainly, there appear to be no osteological or myological differences between these 

Scaphiodon macmahoni was included by both Berg (1949) and Mirza (1969) in the 
synonymy of Cyprinion microphthalmum. However, I find that S. macmahoni possesses the 
same mouth morphology and degree of dorsal fin spine serration as does Cyprinion watsoni 
and thus it is included in the synonymy of that species. 
DISTRIBUTION. West Pakistan, Iran and Afghanistan, and the Helmand basin of Baluchistan. 

The taxonomic status of Semiplotus dayi Fowler, 1958 

Fowler (1958) considered that Scaphiodon aculeatus of Day (1880) was a misidentification 
and that the specimens concerned represented a new species of Semiplotus for which Fowler 
(1958) created the species S. dayi. 

Although Day's specimens can no longer be traced, his description (1880) gives no cause 
to think that he was not describing Scaphiodon aculeatus. Since Scaphiodon aculeatus (Val. 
1844) is a synonym oiCapoeta capoeta (see Karaman, 1969) it follows that Semiplotus dayi 
must also become a synonym of that taxon. 

Relationship of the genus Cyprinion 

In this study the presence of a sector mouth in various cyprinid taxa is seen both as a 
character indicating relationship (synapomorphy) and as one independently evolved as a 
parallelism. Taki (1975) used mouth morphology, degree of dorsal fin spine serration and 
ossification, and the presence or absence of barbels as the criteria for determining relation- 
ships amongst 'semiplotine' genera. Taki considered Onychostoma closely related to 
'Semiplotus', "Scaphiodonichthys", Scaphiodon and Scaphiognathops. On the basis of these 
supposed relationships he recognised an Onychostoma group and constructed an elaborate 
dispersal hypothesis to explain the distribution of the included genera. 

334 G. J. HOWES 

Taki (1975) supposed that the onychostomine genera were '. . . derived from Barbus- 
Varicorhinus stock'. As he indicated the group's 'origin' to have been in northern Asia, I take 
it that he was referring to Euroasiatic Barbus and that his ' Varicorhinus' was in fact Capoeta 
(see Karaman, 1969). Taki further confuses the issue by referring both the middle Asian and 
western Indian species to Scaphiodon thereby implying their monophyly. In fact, Taki's 
middle Asian 'Scaphiodon' are all Cyprinion species and those along the western Ghats of 
India are Osteocheilus (see Hora, 1942, for synonymies). 

The remaining genus included in Taki's Onychostoma group is Scaphiognathops. 
Osteological data on this taxon are completely lacking; examination of the type specimen of 
S. stejnegeri (USNM 90303) reveals a quite different jaw morphology to that of any 
Cyprinion species (see figures in Smith, 1931 and Taki, 1974), or for that matter, from any 
Onychostoma species. 

Taki's (1975) hypothesis of relationships and dispersal are falsified by the conclusions of 
this study, namely: 1. That Semiplotus, Scaphiodonichthys and Scaphiodon (part) are all 
synonymous with Cyprinion; 2. That there are no synapomorphies linking Cyprinion with 
Onychostoma or with Capoeta. 

The relationships of Cyprinion are still in doubt and the sister-group remains unidentified. 
This is due to lack of comparative anatomical data for the corpus of species currently 
referred to the genus Barbus. Initial researches cast doubt on the assumed monophyly of 
Barbus and also indicate one group within the complex which could be the sister lineage to 
Cyprinion. The taxa involved are the north African species B. luteus, B. paytoni, B. callensis, 
B. waldoi, B. rothschildi and B. sharpyei. In particular 'Barbus' paytoni possesses cranial and 
jaw features similar to those of Cyprinion viz. deep and ventrally curved -ethmoid bloc, 
transverse dilatator fossa, an almost identical posterior neurocranial morphology, broad and 
deflected labial surface on the dentary, and a broad anguloarticular facet. The shape of the 
quadrate, the hypertrophy of the interhyal, and the inward curvature of the interoperculum 
appear to be further indications of relationship. More research is however, necessary to test 
the hypothesis that the north African ' Barbus' species group is itself monophyletic. 


I wish to express deep gratitude to Drs Humphry Greenwood and Keith Banister. To the 
former for his critical reading of the manuscript and the many helpful suggestions that have 
made for its improvement, and to the latter for spending so many hours in discussing the 
vagaries of the carps. 

My thanks are due to my former colleague Margaret Clarke for her assistance in compiling 
the species synopsis, to Robert Travers for preparing alizarin specimens and Gina Sandford 
for typing the manuscript. 

It is a pleasure also to thank Dr George Lauder for our many stimulating discussions on 
cyprinoid myology and functional anatomy. 

Special thanks go to Dr Christine Karrer for translating, from the Russian, passages of L. S. 
Berg's 'Freshwater Fishes of Iran'. 

Finally, my thanks are due to Dr Richard Vari for his assistance in locating H. M. Smith's 
specimens in the Smithsonian Institution and for his hospitality during my visit there. 


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8 (4) : 783-858. 
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species ofScaphiodon Heckel. Rec. Ind. Mus. 54 (1) : 1-10. 

& Gupta, J. C. 1940. On a collection offish from Kalimpong Duars and Siliguri Terai, Northern 

Bengal. J. roy. Asiat. Soc. Bengal 6 (2) : 77-83. 
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Cynodontini and Characini. Bull. Br. Mm. nat. Hist. (Zool.) 29 (4) : 201-248. 
1978. The anatomy and relationships of the cyprinid fish Luciobrama macrocephalus (Lacepede). 

Bull. Br. Mus. nat. Hist. (Zool.) 34 (1) : 1-64. 
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the subfamily Cultrinae. Bull. Br. Mus. nat. Hist. (Zool.) 36 (3) : 147-200. 
1980. The anatomy, phylogeny and classification of bariliine cyprinid fishes. Bull. Br. Mus. nat. 

Hist. (Zool.) 37 (3): 129-198. 

1981. Anatomy and phylogeny of the Chinese major carps Ctenopharyngodon Steind., 1866 and 

Hypophthalmichthys Blkr, 1860. Bull. Br. Mus. nat. Hist. (Zool.) 41 (1) : 1-52. 
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asiatischen Arten des Genus Capoeta (Varicorhinus, partim). Mitt. Hamburg Zool. Mus. Inst. 

66: 17-54. 
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ofcichlid fishes. Amer. Zool. 20 : 295-314. 
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Cypriniformes). Bijdr. Dierk. 33 : 3-35. 
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Manuscript accepted for publication 23 September 198 1 

East African Cichlid Fishes 

The haplochromine fishes of the east African lakes 
P. H. Greenwood, British Museum (Natural History) 

May 1981, 840 pp, 3 plates, 350 figures Hardback Price 25.00 

This volume brings together, for the first time, Dr Greenwood's various papers (some 
now out of print) on the taxonomy and biology of the species of haplochromine fishes 
from Lake Victoria, east Africa. Also reprinted are his papers on the haplochromines 
from Lakes Nabugabo, George and Turkana (Rudolph), two papers dealing with 
the classification of the genus Haplochromis, and a recent essay on the explosive 
evolution of cichlid fishes in Africa. 

An index to the 200 species dealt with in this book (and their current generic place- 
ment), and a general introduction to the evolutionary and taxonomic problems posed 
by these biologically complex and fascinating fishes, are included. 

Published jointly by British Museum (Natural History) 
(exclusive U.K. rights) and Kraus-Thomson Organization Ltd, 
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Titles to be published in Volume 42 

The tick collection (Acarina: Ixodoidea) of 

the Hon. Nathaniel Charles Rothschild deposited in the Nuttall and 

general collections of the British Museum (Natural History). 

By James E. Keirans 

Hydroids and medusae of the family Campanulariidae recorded 
from the eastern North Atlantic, with a world synopsis of genera. 
By P. F. S. Cornelius 


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